Difference between revisions of "Journal of Morphology 28 (1916-17)"

From Embryology
m
 
Line 44: Line 44:
 
Ethel Browne Harvey. A review of the chromosome numbers in the Metazoa. Part 1 1
 
Ethel Browne Harvey. A review of the chromosome numbers in the Metazoa. Part 1 1
  
C. M. Child. Experimental control and modification of larval develop ment in the sea urchin in relation to the axial gradients. Eight plates. . 65 Roy L. Moodie. On the sinus paranasals of two early tertiary mammals. Five figures 135
+
C. M. Child. Experimental control and modification of larval development in the sea urchin in relation to the axial gradients. Eight plates. 65  
 +
 
 +
Roy L. Moodie. On the sinus paranasals of two early tertiary mammals. Five figures 135
  
 
F. X. Williams. Photogenic organs and embryology of Lampyrids. Ten plates 145
 
F. X. Williams. Photogenic organs and embryology of Lampyrids. Ten plates 145

Latest revision as of 20:50, 21 May 2020

Embryology - 7 Jun 2020    Facebook link Pinterest link Twitter link  Expand to Translate  
Google Translate - select your language from the list shown below (this will open a new external page)

العربية | català | 中文 | 中國傳統的 | français | Deutsche | עִברִית | हिंदी | bahasa Indonesia | italiano | 日本語 | 한국어 | မြန်မာ | Pilipino | Polskie | português | ਪੰਜਾਬੀ ਦੇ | Română | русский | Español | Swahili | Svensk | ไทย | Türkçe | اردو | ייִדיש | Tiếng Việt    These external translations are automated and may not be accurate. (More? About Translations)

A personal message from Dr Mark Hill (May 2020)  
Mark Hill.jpg
I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!
Historic Disclaimer - information about historic embryology pages 
Mark Hill.jpg
Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Journal of Morphology 28 (1916-17)

J Morphol. : 1 - 1887 | 2 - 1888-89 | 3 - 1889 | 4 - 1890 | 5 - 1891 | 6 - 1892 | 7 - 1892 | 8 - 1893 | 9 - 1894 | 10 - 1895 | 11 - 1895 | 12 - 1896 | 13 - 1897 | 14 - 1897-98 | 15 - 1898 | 16 - 1899-1900 | 17 - 1901 | 18 - 1903 | 19 - 1908 | 20 - 1909 | 21 - 1910 | 22 - 1911 | 23 - 1912 | 24 - 1913 | 25 - 1914 | 26 - 1915 | 27 - 1916 | 28 - 1916-17 | 29 - 1917 | 30 - 1917-18 | 31 - 1918 | 32 - 1918 | 33 - 1919-20 | 34 - 1920 | 35 - 1921 | 36 - 1921-22 | 40 - 1928 | 47 - 1929 | 51 - 1931 | 52 - 1931 |
Historic Journals: Amer. J Anat. | Am J Pathol. | Anat. Rec. | J Morphol. | J Anat. | J Comp. Neurol. | Johns Hopkins Med. J | Ref. Handb. Med. Sci. | J Exp. Zool.
Links: Historic Journals | Historic Embryology Papers


Founded by C. 0. Whitman


edited BY J. S. KINGSLEY

University of Illinois Urbana, 111.

WITH THE COLLABORATION OF

Gary N. Calkins Edwin G. Conklin C. E. McClung

Columbia University Princeton University University of Pennsylvania

W. M. Wheeler William Patten

Bussey Institution, Harvard University Dartmouth College


VOLUME 28 1916-1917


THE WISTAR INSTITUTE OF ANATOMY AND BIOLOGY PHILADELPHIA


COMPOSED AND PRINTED AT THE WAVERLY PRESS

Bt the Williams & Wilkins Compamt

Baltimore, Md., U. S A.


Contents

No. 1. DECEMBER

Ethel Browne Harvey. A review of the chromosome numbers in the Metazoa. Part 1 1

C. M. Child. Experimental control and modification of larval development in the sea urchin in relation to the axial gradients. Eight plates. 65

Roy L. Moodie. On the sinus paranasals of two early tertiary mammals. Five figures 135

F. X. Williams. Photogenic organs and embryology of Lampyrids. Ten plates 145

E. A. Baumgartner. The development of the hypophysis in reptiles. Sixty-eight figures 209

Fernandas Payne. A study of the germ cells of Gryllotalpa borealis and Gryllotalpa vulgaris. Five text figures and four plates 287

No. 2. MARCH

E. R. Hoskins. On the development of the digitiform gland and the postvalvular segment of the intestine in Squalus acanthias. Thirty figures. . 329

Walter E. Camp. The development of the suprapeiicardial (postbranchial, ultimobranchial) body in Squalus acanthias. Twenty-nine figures (two plates) 369

Grace A. Wells. The skull of Acanthias vulgaris. Twenty-two figures ' (three plates) 417

E. Eleanor Carothers. The segregation and recombination of homologous chromosomes as found in two genera of Acrididae (Orthoptera). Five text figures and fourteen plates 445

P. W. Whiting. The chromosomes of the common house mosquito, Culex pipiens L. Seven text figures and seven plates 523

Robert T. Hance. The somatic mitoses of the mosquito Culex pipiens. Two plates 579

George T. Hargitt. Germ cells of coelenterates. III. Aglantha digitalis IV. Hybocodon prolifer. Fifty-five figures (six plates) 593

D. B. Casteel. Cytoplasmic inclusions in male germ cells of the fowl tick, Argas miniatus, and histogenesis of the spermatozoon. One text figure and eight plates 643


A REVIEW OF THE CHROMOSOME NUMBERS IN THE METAZOA

Part I

ETHEL BROWNE HARVEY

Cornell University Medical School

The following tabulation comprises the chromosome work that has been done since the pioneer researches of Van Beneden in 1883 until the end of 1915. The list is as complete as possible, and recourse has been had in all cases to the original publications. All the authorities have been given for each species, even when the accounts are conflicting, and these are given in the order of priority. All the species are classified and the various phyla, genera, families and species are arranged in alphabetical order. The present paper includes the Annelida, Arthropoda and Coelenterata, and will be followed by a second paper including a tabulation of the other invertebrates (excluding Protozoa) and the vertebrates together with some general remarks. Some of the most important abbreviations are :

chrom = chromosome pa = parthenogenetic

cl = cleavage p.b. = polar body

div = division . prim = primordial

el = elements pron = pronucleus

fert = fertilized som = somatic

mat = maturation spc = spermatocyte

oog = oogonia spg = spermatogonia

"X to pole" means that X passes undivided to one pole. "XY to poles" means that X and Y pass to opposite poles. "From correction in" means that the tabulation as given is a correction of an earlier account.


JOURNAL OP MORPHOLOGY, VOL. 28, NO. 1 DECEMBER, 1916


ETHEL BROWNE HARVEY


<


z

O H O

o h a

> w z




t- &



<! g


00 23




^ O



"o a


dc


Bo,: o2« 


s"3 ft

3 *




flre


"E ft03


DQ&i .






3g^




£ <j


0"fa




o




o .







- s







2S Jg.S





c3 03



0> OJ



43 43



02 03


02 ft,






§82


o*


Z


■b§


•*


OT3





■>;







2




O




K




<




e


Ma?




0j„O


o



'bg


•>*



O-O







-— . -~~










bc2 w-2







o o oj o




Pq-C^TS


QC Tf



oooo




CN—KM"-"




(1 II




V




"bo*







o o




oo




3














E2




o ft



H

£


u


M


js a



3


■a .3




-°'E




.g'3



o







.2 o


Ed:




■y


•£


OB


a



m




ci"


Oj CN


J^




oi




-T



-f









Cl


c






a


i 3 <



b



jg


&


f


^"3


a


02


d


S




-v


d us


00

o

o


d


Od


O . O ft


B

3


a"


3



35


Cl




43


c










43



?>


. a




. Morp r. Jou 199

i. Zellf


B


9 M

r3


js-°

-° B


So

^fi -So?


4^


J 3" 2&ft


a


3 B



3


^c*




H


o


.-=



~


• U


-3 .




O





3


5


ag


"mPh


mPh


Tcfti


4!


3


2


-1





>-s


<!


■3


W



IVl


M


"3



1-5





_





T3


O









3


OJ






3


P'









43


rQ






B












O


p



p



CO

O


-a

3





p





02


w




S>,


s^


^>>".


B


p




•^




03


T3

3 B


T3


CS


M >



4*


>



.


.S


CO



44 >





o



j5


O


ova


3 a


4^




O




o

o


co o ^h .-, o- o- o



•'a?


y


•ll


B



T3

'0




fepH


fe


>



>


>


>


« 


O


~,



>


fi

o


a


o









■b


■b














CO






c


-S










rt


rt





S~


o
















K


ca

















2
















O

















>-*
















■O



w









1) CO


■b


■b














1



CO


CO






























^>
































■"" l















































«i















M<








CW





Of



ex

.CO


'b


■b












CM


CO


1




CD



CO










CO



















5
























M




te



tf tn


M tr





U


Ml









c



ft2


flO





c


a






O




o



to O


10





OQ







(M




■*



<N (M


■*>*





C^


CM






CI




CI



CO 00


COCO





CO


CO











u









B

























B

2




3



-*>

3


C





B



.2

'(3


'S



3 >

B




B




-a

03



E

3


O

s

M


c '0


c


1


l-t



a

d



C3 00

3




'






,



'S


'E


E

4:



43


43




3






43



J3 O


T3



2



£


s



0)







a



a


'E


>i


3



3


3



S




<




UJ



a


£



n



iJ



J


►J



S 1


CHROMOSOME NUMBERS IN THE METAZOA


« 


to


&




jg


^J


!S


c ~





ft


© •


a <


Oft


iC







AJ


H"


©



8

.3

3





.a Ml


cc


J2 M




J" 1



o


Hofri


2 ftbtP-i




<D


H


1-1




-a






a


r~



o


c3fiO









«>>


o>


o


^


M^iM







>


> 8



>>

.3

c3



0>a


o.


T3


r ^


T3




>


>


>


o



o





CO





o









co







o





o


O


o



















o


bO O O




o


CO


•*

CO



c5


3


a






©









©




H



d



3


3




M






3




3



a



>


>>




u


j3


.3





O








at


>i



J3


©


J3




S


« 



Eh









^ :3




03:3




Ol^.

















£


Ott










CO .3


T*


r ji


<o a




cm ©



• H5





c3-2___

._. CO 3



8^


° III








§io



n


.■S»-c


gio




N


N


s


N



CO




CO











CO






3



o


a




3




a




cj


c


CS




a



3 O


a










&


PC


3


ft




a




a




©




©




a


>


s


H





















ft


o


'Vi



o














■a:


3
























^













K*






w


o







e








■bo


b



bo




ii*




■*•*




bo


■b


O


bo




■*-<*


03


-n •*




M


tr M


bO




ft


C.° — „


ft










00


00 CO GO CO


00













3


3





















CO


















o


c






C3

a



ft

CO I


ft

ci

ft




3



3 h


3




























O


r


o


O
















ci



cS


2




XT



a


CO


CO





£


"3 §





u~



o> a 01 "





o





CO



o _J



r- o

U0 CM


ft


N _r





■^


"Sg



dft

co cm*


CD CM


-2 a





ft

C5i"


ft

CM


a ft

o ©?>•

CO





§«5 m

©

j3 © "i





g'


3


^c O ©



3 O

-5

ft




s


o


g^Sa



n<


> N





<


J


N<J










i











Q




t~°°



OS Cp





p


T3


o —



p?








3




© ©

© © 3 3

a S o o

mm


US


"© "©

-3 J3 o ©

CO CO

O O





CO

©

3

'53 u


03

© o

'3 -T M 3 © O


© ©* 3 3

- Kh

o o

J3J3 © ©



3 >-5


M M





CO


o 4 "


QQ











co

© "o a

o











b





















ai


c3



f>


o







"c5


3



•*


CM







a


09











M










c^


w











-^'




o





b


b


b

m u

0.





CM





■>*


•*





















Oi











b













o


O


o





bo


b


CO

B



■*


-f


CM





•^-^


<*


C8










a











m a







©


©




ag



o


tad bO 8



3


_r:


M tt 3


M a


CO m





ftO o



c


"3


ago


ft O


'm fo



CO



CO O co


O


yj



co O co


CO CO


B.H



A



-f-^-V


"*



1


00 00 oo


00 00


■3 s a a



CM





00


J




i<-*





m


CO





oi


.2


.&
















o


M





©


t->


©





"c


3





3


3


3



ci


9


ft





ft


ft


a



s


03





03


C3


a



03



A





.3


X.


J3



.0



©





O


©


©





o





O


o


O



a



M





R



tH











+^>




©


o





O


o


O



CO


a


>>





>,


>.


>.




>,


H





u


u


M



s



J3





£


JZ


J=



©


C8


a





o.


a


a



2



s


o





O


O


O


ETHEL BROWNE HARVEY




?





S





DO




a






>




■a" 35




a


. d


cm



5o


gCM

ao °°







a


.a


<d


PQ



x. c



J3



£U


•s*

b



«.


P3


-3


K



■**


co






>





a a


~


03







a o


c3 C


%


o



C



J3



41


£



to





M





K





■<





S





m





a





a









T





SI





e




»




u

1





a





z





cm





H





>• U





o


o


b


1


10


■>*


CJ






00









6





Z





= 22

o h a - - *

o ^ o


be O

o



be

a


o



00








a





z





<!







co


. £




s


B








o











to

a


03 9


'3


00

'S


Ed


a


o


o


CO


to


CO







3








a


a


a



o


O


o



S


s


S



o


o




H


H


H


d




'c

V.


BO CN



OC


d


■SO


.2 cm a.

Eco


35


O "^
















3


<


a



d

.a


"o o N


0) >

a


■ a a

'5 03 X

> C5


O >

>


s >


p








a a S


d

W


"3 o


lis


OJ


(goo

^ CO

ro


3 o


OS



"5




C


32

o

•-3


lid



bis S.


■gSd


>-=




« 




<


O <



<


CO

c




C5



















— '




c




12




CO


CO


r _


r ~



CO

p


cj







Ci


o



C g




V.




3






c




1?


£




£




■o


C5


g


I J



_CJ




o




03


2


o


"a


•-5




X




a


3


Q


£ n



m











~


&



c

^'









rt




cc


d









C


5




rt












n















cj









a


03


-d

=


o


OS

a








oj


»


ON


ot


IS


)0








K


2



X


£


■a








X


a


a


3


a








o,ai



PfO


03








cm




CM






































O




o



'b




an





C5




o




2 ■*



bo

O










■b


"bo




























o




e>




o


o


.

.- .






o




c


03














■b


bo



























O


o



o




o


ex




-f





c




OS


es


Cj. co


'oi at

oc 00



8











^_













tl













— ?





a




S3









c


£












00 Ci



cc



.—


h £




e









CO


Si oc 00










C


a '■














o


o •














£


S :














03


C3 ■




=!











bO

tH

<D C.


u

o ■ & :


b (


1


3













>>



u








CO


oo ;












3


3 ■



Q.




a







u


kl •





C3








CJ ot


eg co



^rt











■« 3


o



O








c.5


— o


o




_g








O a


o o








_s




-> c

■s u





3


C E


,






1"


§ 5


'c


X




<




«<




O


O




'/'.



X




d

s

c H



8




73

1


>>

X


£ p aa

bo


a

p, p,

<









B

3

o3 | O

N

s


CHROMOSOME NUMBERS IN THE METAZOA




r ec


a


rt


&

o




CtN .



-csi r


a






JB*


CM


aw •





o „ X



a gws N


sis


-a



£££





a





C-lhJ


« 


o






o


o


t~


S3






o


OK>


+


CCtcM











o C



■**< CM



CM -H



ce



03










m a)


o


aa


a


o3 4


05


as


S


0) o


a


03 C3


03




o3 03


03


J3J5


J5


h


H


H







0(M



£;


r~






a**



a


a


x .





»>



CN


<M






^ .



£>


X>


3 s





^



C3


.


xj


>-5



•a g





£ 6lD


fi


O


<!



so


N




CO


CO


p




o


13



-1


-c






S3



3


3










CJ



91


o


o



O


o





a





03









£









o

























03





> CU

















XI O





o


o





o









o




o


o




o














rQ .~,





o













II a














s




° >





o









.— *





.J »











-C ?3















o ?




3


M >




73


03 03




3 3





a c





o O




3


o o




3


o o





o o




a





3


J3J3





Cl,0h




X
















t~


eq







a


a








o






^



.



















N



N


co





U




-d










h-l


<



<














o


C



a



CD



a


>>


2



a



■o



o










o


1-5



i-s


Of


O



o

o


00



CO







> k f









o














o






w







-i









a

03



E


Si




a


d




S o


3




s


>


rt




a




_03



-fi.O



0)


o a


5


O


£



a.



s ^


< Q O Ph

o








o



3


a



fe








GO —




Anz

Soc. 7, P




8 '


N -<


a>


o




-a




2 o.

M P


? ^"



-3 -w


u 3



^ tf



"



S



o



>>





03



II



i3 d







aa



"bot


CM CM


b


-« 






1 --^



4>T3


M £


a o 3




oo




g ~£







■cH





S



3



d







B




=



>


00



'S


~


a






i


T


'-3



a



d.




CT C




o o




o

- *

00 CO




C^ Oi (M





a • a

. a

2 -t



a

oo

CO


"-•OO^H






r - »


X


XJ



J3








r


1-5



J3 03







fl°°


S


8



8


<:«« 


N


N



N


0«5 0)






OOO











ODD













•*



















s:^&


(

h









«PQ« 


c


a



a






03


i-MJ


Ph


Ph



Ph


. a



i






DQ














= 2




p



<D ?



a



a







o

a



CD


s



sgs



aa


a


Xfa.S





a



CM



M



X


■b
















CM






•O






CM








■b






+






t~










■b





"5




'b












to






a






CO
















ua











03







T3





m



a





>



fl









C


^,










>





o

03


a.


a



03

a

CD


a


c


3




g


P



03


a


c


2



-a



a



a



"5


n a



>,



e



a


<



<


<:



-4


ETHEL BROWNE HARVEY



OJ




o>


CT:







~ Oi





1^


OS



o




Q





O O




CO




"5



U3


IC


a


■e


m


IO LO



Tl


"5



d



d


d


H

d


. CM


d d


d


d


a






d



03« 







o


00



X


cc



cc


^


CO CO


\


10


X


z


CO


0.


eo


BQ



CO


CO CO




CM



a a


J2


M


J2


•Q


-C



£1 J3


1 ""

ji


.a


a


H



u.


H



M


t- f-t


cT



h>


fa

a


J3 C3


_r


— 0)


"5


"5


"3


_C3


"3 "3


!s _3


CO


-a

03


a


1-5


~


>-»


>-5


Ha


>T> >-5


"0 "c!


"J


►0




O






olc



O O





O


a


O



O


£ fe


O O





O



o


oj


5


O



3


O O


C3



O



N


►j


N


N


5


S)


N N


►J ^


N


esa









10





r^










p





p



a














a








>>





>.



> K a





'*



■^


u


■>J" ■>s<



> 1ft



«*


X








CO <





"



=:



s


"


'


s



n


u


>>


hi


hi





u C





c


a


o




<0


>>



to



tc


c


o


a


C


c


a


hi


a


c


s s


a a


c


a









c







'3


OS


■3


'3


« 


CO


'3 '3


C3 03


'3



Sh


O


&,


ft,


O


ft


S


£-c Oh


O O


S


Ph






_e




»i


3













E «a


03






00

M


a



a


c p,


fl


^


U 3^






a


< S

a




J2

z


X


I "3


"3


«•[edit] "-1 •-



Q






~



■b







fc








CO







a














>H



b








5 '




o


b


CO


b


b


b


r o


b


b r r



CJ


»


b




1


cm


CO


CM



CO


i£> »-i


J



CM





CM


C9







CO




Fh



CM












DO



























C














55














O go









bO u






- - _

SSi|

< s





bl



M


CO










ft

IT. CO



D. 00


00 O + +




"3



«,a





O)



Cl


cocs






o °









CM <N






<!


































EJ























3























M























_o



00

5


JO

'c





C



3

a a


d


T

a



£



e


?



9




a

-0 'I


3

31

3


o

3

a. a


a

x>

B




a> _=

T


c ■a


0,

O

"o


E S



'd


c


d )3 S "3 ■> I

■5 ^

>

2 >>


a

_; 'Z


00



EJ

a


CO

3



3




El



r "0


a


ga

S 8


CO

O >>



= 9 b


03

e

00


j3

J3


00

>>





"_





a


l.





J




O



Fh


~



h


X









CO



01


C


C.


22



cT


ft




^;













3


N


N








j:


-=


O






^i



<


<


Hi







00








O





a


^.


^






CO


CO


O



b




O




T




00




^







^^>


b



00



■*,a


CM



S^


CM


















cC



a


D,



CM


CO



3




O







3





3


pq



































O,


3







3


00












3









3



.2-S


3


3




H


r.



CHROMOSOME NUMBERS IN THE METAZOA


«a>





CO



Oi


»o



«o



lo



Ci




Oi


CO


S







CO i



CO


io




•^




■w




Ci


CO








CM





^*< CI co


5


c c Eh


■«•


s ft

Q.U5

CM«* 00^3 . a> »N c ^;


T3


■*


CO



S

a <i

goo


Zellf., 13, p.

Jahrb., 5, p. 2 mikr. Anat. 00


ftio ■** r

r"3 <5ja


a

co" "a5


is


CO

H ft "3


Zellf., 3 p.

Anz., 34, p. 4 Zellf., 5, p. 3


ft

CO

r ft


3

i W

o

a o

o^ H a


ft

CO

3

ft "3

N






3 .



o

u.


3


pC


-fi



s


«J3


-■>


JT -C


. ( o tsj


W


J3




4j


a




S ft 3


1


<


1


1


Q



a <


O o O (h


"c


3 a>

-5


u




to



ifi



















Oi



r




e


o




=




Oi o




OlO


CM



o



3



U


(MiO

Oi OS


33


^



j.



a

^J<



C5


MM


m


Oi


M





Si



c


o






c


o o


t


o


o o


u


o


o





(



tt


o







a; a>


c



a> a>


Oi






MM


cd o


u o

hfl


c3 *



t


o o to to


£


g



a


c 3


'o



o

>



E


O


o




3


+


5



a


2



03


Si





C


o3ic3

02


IS


fH


E.


s



"S 03


J


03 03


J3


cS




£


WW


l^kl^,


M


t^


a



K


ss


c


m


SS


o


(S


s












o!


a>





















5





















3




















+ °





















a





















o 1


CO



CO












~c





a


-z


-o



-a


T3



^


■a








eS







T


ca



03


03



03


03








t






o


t


H



IH


s




kl

































a


a;



a>


CO



CO


CO












~


•2



































C





CO



s


5



b


Q



b


Q



Ot



"oi

















03


to




II



o














o





















a





















CD






o

eo
















O









































3


■a!




J[


















cc




O

















II






o






o













3

















o

cc




c3 * CM




















CO


















"o




"a3




3

2


1













"*








3














CO




"*



o


o

3


c


c*



o



o


o



o


t>



II




Jl



CM



. t^




t^



Oi


Oi



Oi


Oi

























o





o


o












o



ae


o




















o


















~ l





















-r





3





































s


a



M 3








£




s




S




o


o



O


o








c




o




o







C


2,-3











CO




CO





-t.



CM









T



CO




CO




co


o




s

















1-1


o




°




















>>'



>>



>>


a











3


CO

3




3 o

a;




ft

03



ft

03



ft

cc)

CC








3 +^

03


3

"3


3 '

03


+3

03

'a


'a





tt


CO



to




CO




Cf


CO



-a


."5


2


3 a


3 C



p,




3



3



3


3


tr




3



'a


'a


'a


.2 c


o d



c3




ft



a



• a


a


'c



X


!2



3


3


3


-° s


3 S

CD

13



h



c.


a

a



S

03



a


S

03 »


SE



2


S


IS


_o


o


.2


a





b




03 °

a



a


"d


e


3


IS


-Q


x


o



C




o



o



n =


a









to C




s




2 »



O ff


'


h


C


,


z


ft


"z


i &


a


ft


a


a !3



s



E


zS =



■S ~



_c



JZ


_o


c


_o


_o


_o




o




c c



c-.£



fi~


a


t.



"Z


3


c


3


"o



o



c




ofi*—



ci -—



OS


03


>



>


■. >.


>


1 >>


>i


>>


>>



<



c


O



o



O


O


C



C


C



C


O


O


O


O


O



ETHEL BROWNE HARVEY




o ■* ■*


$8"



CO


3



CD CO


OS




3


to

CO


OS


3


g


a


ft


do

ft


ft


O


ft


ft


Pi


d


ao.


o.


ft


ft


ft


s


co


CO


■*""»


eo


CO



o

00


lO


CO


CO





cc •>




pw

















c? ft Eh ~











a































w


tS3


S3


<^N


N


CM


■*


N


KJ


N


CS3


^N


M


N


N


CM



J5


ji


— :js


ja


-C


> 01


J3


,S


rG



fj

X


&


43


J3








B.S












<


<


N<


•3


<!


■3




<


o H



3


■5


3


O CJ


S J2


2 3 £ m § o




■cc


fl







C3


C3 C3


<!


pq


SS


+ a


Q Q Q Q


Q Q Q Q




•n o





cj fl o


•-3 1


_o « jo


o «  SO

o


u o


Ss &

o o

J3 .ft

>> >>

-3 -a




c, ii c. _o _o _o "3 *3 *3 >>>>>>

o o o


.2 >>~

•3J3§


"3.S 5




o o o o o


CHROMOSOME NUMBERS IN THE METAZOA


.a

co 3 ■*



CO

CO

CM .



c -*



co CO



3




CO CC!



0 CM


5


CO


CO


10


K




CD CO


r^


-*•


<d


CO


p, a p,



dus



d


d


d




d


d^


d a


a


C3

S


"b


1 '


p. d a dd


•r


c

. i

c Z




r"3



CY



u


'


CO




« 




co"°"


<

H


<!

,A4



CN


CO


13


CO


P3


55»



























1^






N







N



1*3



s


n So


ws»


1


CD

IS]


t5


CD


a


.c


>


■*; J



— 1 (


.c





ji




X





^"^


§1d


c


^3



j:



"o~


e


£


21 "



0<!



S3



•-j u





"0*0

o o


o o


o


ft2 c

a


iw


c


c


e


O


<


c


<



CS3



<



-5


<




<



NN


NN<1


<


■3 •«!


iJ


<


<


<


<


e





as o










=










c:




o>o






p^






















p— 1



. r




" J"










'


OtM


o^



•&


■* us

caoi







" ." „


A


It


OS



^^



c




as




^



C3i OS



Oi




as


Ji



a*


^!^!


c


1

o



o o



c



c



p




C








»?;


p


C



a


O CD



c


s s



o o

CO en



c





3




« 



CD CD


£ £ £

CD CD CD



CD CD


J3


■ b"

3


9

C



E



CD CD

■ J3J3

II


■s ^



c3 c3



B



■♦3

03


nj




"S


O CD

cS:oi

o o o


o

c3

.•3:3


a


03



S


£


oS rt


X


o m



SS



PC



£



£




s



WW


www


W


«« 


M



<


PC


ss




i

-B


CD

S



0) C3


a


3


1




1

o


s


OT3


CN









CD 03





CD

ss




+


5

Q


ti


o a

O


tl


+ o

CO

CO


3


+ co








3 . I 3





S 2

■sis


r



■§


5


-3J3


■c


2


T


?


-0



a










n'Z.







o o


CD CD


c( a


o



1 o

D


99

eg


a

9



g








T3 CD

fi'S

h 03


c



03 CD CD— c








-3


<->


-3




eg


~


3


'O


o


3


~


4)








■t^cM





3+


6



o



CO



« 



u:



CO












Q


(5



C






















o 5



























^ t^



























ftp.












o







o






'bo


'bo

r




■bo



























CM CM












CO







CO










TT











































































CD


CD















o












00


00





o










t^












II


^









































co












o


O














































































CD


CD

















,










CO


CO




o








o


o



o



o


o


O




O


~*


•"*




CM


o oo




'


o



r^





CO



o







£


||




T



-*-d<




W3


eta




















^



t-H ^.





IO

















O


o


























00


00















Eh












^_^


^^















co












CD


CD













a



1



c





S






3

W>3


3

U)3




'B


ft £




£





o





C






CO






S o


S o




CM


3


O



~


c 3




X


CO








<*






00 "^





CM


e




CM








'3










































c











w












o


CD





a





















0C


CO













CD





























f.


i


5

ej



3

C3



a



z



3




|





3




3


Z


3


=


3


.2




Ej


1 c £


o

-a

a



En



c



E



J2







3




3


=


3



5


-?



-is c

3



"3 a




E C


<



e


i


3 H O




z u u a



a


C

CD

u

CO




B CD


a


B CD


C

a. j D


C a

a


>


B u


>


OE


cc


ea



CO



















ac





c


. C


. ft



a



c


i


£


i


ft




C


4


a





ft


I


. a.


C


. C


, c


. ft


_c


_C


jg



_o



_c



C



_c





o


j2




_o


_c



_c
































c


5


"3



3




^C






c



'u


"o




"o


c


CD


c.


G



CD


>


. >


> >>



>>





>



>.




>



>>


>>




>.


>


. >.


>


>


>


>>


C


u


O



O





CJ



O




O



o



o





c



C


O



Cj


c


O


U



10


ETHEL BROWNE HARVEY



to



d


t^



CO



CO




CO

3


s



9



8


1 9g



J



t~r


OJ


3


CO




co


T)<


3


•a

3



•a

3



T3

3


a





0.



a


o.


a,


a


O.


IB


CO



W>



W


^2 a d


o

z



^j


O


r^


w


CO


>o


"5


CO


O




O



O


S ■co"'


w


1





.Q







C


c



a



a


3 O.

















oj . r*i


w

fa H H


n


a 6


m



c

<


"3


c

■<



N


O— i

Eh



d



1 Eh


d


H d


—"CO)



J3







J3






>



>



b-r



o


P.


"o^


a


So

o




o


y


•32


a



a



'3 iS


tun > ° h



<;



ea


tS3


tSJ


<


ts


<


-<


&


B



B



B


B B



B B


1


03


g


cS J3


M


S


<


O


-a o o u ax: co co

cc) o -w +* HH CO C3 C3


ui c .s

2 §"" ° a ■2 c23 o

■slsll

O C3T3 >


M tl c to ago o


o o O


•>§


c 1O cu


'P 8


CHROMOSOME NUMBERS IN THE METAZOA


11


o

p.


g

o


c


>*

d


00

0.



OS

a


oa

a


CM CO

■»*eo

da


eo

CM

d


O

O CM S) ■*

.2d


<*

d


eo

a


C^l CO

■*co

ad



> "3


CO*


OS OS CS cs

CO CO

d a



eo

d


CM CO

•*co

do.


c o s:


Sftd


CO


-3

a



■*


ifi*

CO



CO


o"


CO fc


CO


J3«-i « 

C5 .■>*

3 ft m


CO


US


CO



d


"9


CO CO



>* cd"


eo .


^3

CD


.i"us



3











3 - r







d


3







3




~


x







N



es

H


<D GO N










CO

w


N —


CD 00 K1G3


CO

tS)




«3


13



IS)


'3 si


3

<


CD

SI


.cc r<!


3

<!


o S


^N




CO


N) SI



(3 CD


13


— a cd




c





A




-a


ta



3 * —



-3





— 03


ja't


J — '



B



jd

M


CD_-J3


o


t.


C5


o


o



cd


d eg Ha


O


CD


o w o


o


U


"b o




Eft


CD CD



"b CD


t

n o cd


1


cu


O


o



h


o


tH


N


o


H


O u



I* fc.



3


tS^


CD


>



N


N



•?


t$<


o


> N


n<;


N<l



<


<5 <!



<


>


N-<



4.











^















^




a











C3









10 lO






cd




3



en







O



g OS


C320





1C





CSO


a


OSO




p






o





o


p



p— <









O— i


O— i



-3


p



J-"




eo




~r


«_oo







p


"


C m



~J


"


"


-d


00- " o . .



C



.sj





o


-M-*



6" -*


MM


-M^<!









MM


c


■ MM



cd








o




o3.ro


Q


o


e? u








CD CD


03


.T CD CD




-M


a>






CD


o





CD



CD CD



S3


09






CD CD



-i -- ■

03

a a <


- '■-MM


-3

a


a a



cd

a


u

a M


-J3.J3


o M


fci cd,


-3 O


o


J3-C O CD



s


a a




•CJ3

CD CD CO CO


^ CD J3 J3






a


o


-tJ




CD CD -H>







M U



CD



CD


35 cd cd


7i

',


S3


'H



red


cd


o3


S3


0J CO 03

03


o3




s


o


o o



3

'Stel


rt

n



a


M



<


w


as


« 


« s


ss


ss



CO


>


M M



fi


ss


n


ss




















CD




4j" o


























03





























CD




3rS


























3-a




— 03 eo<~


























a

C0 N




DO I*


o


















■o


•§



t j



S i=




I'ft


oj

be















cd




03


o3



d






3


















c*


t


u
































CD CD


fl















3)




CD


o



o



°!




G C

CO


o

CD















s




(3


5



p



■*










a c







a a























o o

a a






22 aa













'bo


o




























■*■«"


CM










'bo






'bo














^










r-~ t^







■*•*






































































^^

























































CD





























-*




























'bo


CNI



. 'b


























00 00


II o

CNI



CS1



















































































CD














.
















00











o








o



o


o


o



o



"bo


T*


o bo



o


o


o




TT




»





b


eo


t^



CJ^



1^




oooo


II





eo


cc



eo






















o










































GO


/




























CM


1














M bl




M br


























o-g



ao








o







cd


cd




u









co O o




CO O


o







■*





M b(


r,


oo


eg












GO oooo




-3"*'*'







co





CO O

oooo


c






CO












CNCM








u c



co

3

Cu

3 Fh

eo




CD



co

3 CD


'a

M

C o


'E

o cd

a



_0




'"

a



ed

'S



I



i o:

§

00




cc



cc

^3


1

o



ce +

3



t

cl



'S

a

2



o3



o




o

CD



o a


"3


■a



to


b



_2



1





03


c

-3



o


"o



%
































s


3




3



CO

9


3


3



3


E



3



3



3




s


3


a



CD

a


3



CD

a



a


s




a



a


£


£



£


5



s



s



s




.2


£


c3



o


c



o



o


o




o



o


£


o



c


_o



o



o



o




63





o


o



o
























M


c



h



a ■


a




a



n,


a


ft



a


C



a



Q



a cd


J3


_3


CO



CD


Cj



CD



cd


09




CO



03


03


03



cd


03



03



a



03



o



u.




































5



£




s



'q


S


s



P



a



R



P



5




H



w


w



a



w



w



12


ETHEL BROWNE HARVEY




OS


a




C3




OS




CO


06

CO


9

%


OS

CO <M


ee


a 3°"


CO


OS CO


m


d


d



dd


d


d


d



n


CO


a


CMio


cc


2 „-*


« 


CO


a










a










a

b B

K


"3 pa


id


o

Nft


|n


K


SCOT'S •^ -J

.2 §.2 2





"o


o

H


^•o


.2 2


— c.


.e cj


J3



n


"1


-5"


««J


<


cq n<


«J


<



CO


IQ



COO



CO t^O



"0


K


p


~



5?"



p p— .



^


i>


d


C



a a"



a" a" a"


OS


CD


K


o



p


o o



o o o



H


T3


3


•73 -O



-73 T3T3



3


01


a


03



a a


f

" a a a



c3


PQ


CJ


A


I*


a <x>


0.


CJ a oj


te


4=


O


5



3


OO


=3 C



a

u



cj



CD


o o


tS


o cj o



O



s .


M


CO


ss


3 3S


S


M


03










M










« 










<!










S










W










K










a


"b



b




b




H


00



■*




00




w










H










P










o

1


00



b




OO oooo













55










CM



















H






J.





H










►<






JZ





O


■b


o


■b


o


§ 'be.


o




00


o


CO


00


CO












F









00






o









OO




6










z.










« w cj

2 K- —

<- K H

►j « S

« ™ o a


M o



-f bfi


bl



tJD bC



bO


ao c CO CO



a o

of o

00 00


O O

CO



£8

COCO



a

OQ

CO


















fc










■<;














CO























Is









OQ


~


.2


cj









3


3


"3


3





00





03



.5






3





.2


ua


a



3


>.




1

o


3 H


u

o


a



0.


~c3 3

a


a


'a


0.

go


3 M

s

C3


3 cf

"3

a


3


>>


o

M

CO



X

c t. c c


a

a



3

c

« 

o

03 tC3


CO

c

'^ IE

c

a


i.



2, p. 390


CO

d

CO


ool., 64, p. 2, p. 390

7, p. 462

8, p. 407




Cv


r


s;



oo .



is


i





n3



"a


03 ►-5


1


o tSJ


c"oj



^



CO ^



S -a


^■^a'



t


"5


OS Q

"° O


'3


s g


o o

O u



<


S


N


N


<


N<





CO








r"


00






c



o;







^0J






o o


M


O


^


p





c


c



n" n



3


Ph


i.


a "dis



0,


a



q


"5


ss



2


c

3


"c


2


J5J3



CJ


^


S


1


eg


CJ CJ

COW





!






3



2 X






id

d^


2 o







■J x


ao


X





a egg reduc

a; 1 p

a; 1 ducti


a

of




Ph


p.


Ph


Ph




OS

T3






b


o ^b


O






oooo


a








5








t














DD








o








d













Of <h







OO


00



o


o o


Of _ o 'b



T


<M


CM



» oc


OO



CN












M







c





T




c











'c


a


s a


CJ CJ



-t



P,



1


CO CO



CM


CN




cr






OJ











II

03


cj








J3


J2





CO



U c!


« 




c

a


a



at


03

a





3



O CS


o



£


C



c


s =


s




c


§


a o



SE


§




c


CJ


C

a


fc S.s


s






i

o C


g



T


i


01



1- 1.

-o c




t

T

Q


j

L 5 >


, o



c


i C


- a


C




>


. >


s >>


5


o



C


C


C



C


^


Z


CHROMOSOME NUMBERS IN THE METAZOA


13








IO







id


00





CO


■*<




d


■o

d


c*


d



U3


d




"*.


-"•



>*




"5




X>



X>



M-l














J3

S3

l-B



<


1-5



3


3

pq




_J


X



^



.3


__;





o





o














CS3


<


X


N



«i


pq




kS




•n







-a




e




CO






















a


■*


a


09

Is S3


00


m




s g

c3 P?

H 03

'3.3


p

d

.3

3

3

o £


p H

3

X>

S 03




&


« 


H


is




O




O









c










X>










a










•-< o



to

a
















M 3



a




X























Ph



Jg




■z,


cS










T3


u




























T1









>.


CI









ft.


o









rt








~





4


a




•fa

CO

o





o


S o


O


"h





oo


1

t


"*



00






mS


mS








a o— .










f: '■!> O

oo



CO 00






























3











a











d S







3

"3


J3

3




a o
















"8


a)




~m


<o


X


X


■ i


ft




B 0.


3

a

03


P

r



3

E eg


J3

"3

J3

ft













J3


J3 ft


C


, E


c


ft


o





C3


a


c



"o

ft,


£




« 


C



c


£


a3




»o


00


»•



eo






3



ft




~3

c


t- d




ifl


to


" '.




■»•




03


Cl


t-^



E<


e*^


Lc











"5



m


CO



iO


. eo




Q

CO


o. abs

s




CJ


d




d


ft


xi

2


ft



ft

ft


a . t-*:2

."o3


d




X


- tH




t*J




^


J


<Ji



^


»*J *3


J


DC


X


S3



C


"3




C3


eg


c eg


eg



c!


3 a

a a>


~



s


a


<J s


b


<


ts





s:


o


SI




tSJQ


S3



1-5


a


— .3 O O

O >

d


"5


o





.23 g



OlOr • •** j


d


N


N-<



<


<




a


<JPQ


<



pq


•9JPQ


<




i



































T




J:















Coo

















03

















oi


3

c


CO ■■*«  03 ^P 1



'5

o


CTJ




00

p


~


~


s



S p


~?


r




el

E





O













o


5 eg e> £ 3 3



5





g


a" a*


a"



E


en


a"



c




h


s S




S


o


o


o



o


o o


c



o

s


8-i^



O .IS






^








1


•-^ hh



eg


t



u


u


b


L


1

Z. u


ki



mpq



A


s




<


« 


<



<


« 


<



13



>>>>
















_c



c3^:

s .

id -a


7















'>



c

















3 i


















-3


















a


















3


















w


















e*




o

00

















o

















c


o



o o




O






o






el


•««*



Tf


■*




■^











oo



OC


oo




cc











cv





















£


e












ti





tc


o

















o o













c



o o






u






"c







■*00






•*






e*







ooco




o?


1



00






•*







rt




o^o 1 oot^

















o


















00






















J,


<L :





J^


u














ja


bo >-•





c

3


3














>

03


3m

IB;.

J3-B





>


■3

eg






al


03 3



CE

c


c4

a



03 ft ft 03





03 03

3 3












~











"e


03




03 00



u





r!


e

oa






a


03



r


03



r5


•a

3^3





rt


















c






1


a



£


a



£


£ 03





£


-z'Z





c


a


ffi



i


o



CB


ea « 





Oj


S3 r





. t


M



t

u



u


>^s






>^





<



<





<




<






<





14


ETHEL BROWNE HARVEY





U CM


CO





a


•»!




H o


35, p. 25

. Wiss.B k.Abt.,'


c

<


h#



K

a


cd «S


c

.-3S


5:



b.



. -*j=


eh


s



8


s •


N




•"» •?"•£


.A


a





_ —


— t

Fh




a •


<


-J)



a













>



cm





a

a

a


g a>


a;

3


=:




o 5


3 03


s




§


pq


m


£



X.







M







K







«c







S







a


















"O







V







=







c



















=







o







"_






c c



p



o



o o -


1



CM



aa







'bo*


O






CM CM








tf



_




=


H


a






Eh

o


c


CX



<


1


X


2




Q


a

z


b





cm


c












m


i






H







><







o 1


G

3





Eh

IS


b

o











6







z







q B O





m _



- -


M





9 fc H J K ^



0— .



o o




o v



•* ^




•w



CMh«<




c*e*



1 CM



o o





c*



a





CM



z







•<





















OD



_=



.O



H



3



3



3

H



H



M



p.


a


0}





a


S


3







. a



a
















~








o



o




c


c



c




a


C3



d




t

t*



L.




ca


PS




CQ



'


CO O

cm

"a u o

M

»H

3 O >-a



"3




8. Malacostraca ■ a. Arthrostraca 1. Amphipoda





b

00





'8

u

o o

'm

_s

g 8

J3

o

(• o "3


=3 S <! dS


??3 Pi >-s


d z z z


b b


W _, DD

+5 -2 3

C3 ej 32

s s s

? H 03


< s o


Qa



« 





cc a









"





- 2.




r

=s a


c






' 5



oo r^


£ "2





o « 



O


00





.A



CM


b


c



Hi



~



CC




u



a



aQ



O







to









03


s


C

s>



3


fc


S


z


S



g,



« 


c


s




t- io



a




§


K







n



b


3


5 S





c


'

o

i-j




e






■*


J



c




a


%





u



P

fa



















00





o







a





3





O



<!



b





O




b


oo


b






e










c



3


"o

- 5


s

E


cc


1


"


S


^j


r


E




<


o 1


CHROMOSOME NUMBERS IN THE METAZOA


15





n


CD


00



■ a> ft CO


lO


,J



03 PL,


w





c



CN


a


co"


m


CM


CO


.3 ~

1 S


3

1-5


1


.3


5 8


^ CD 03O o


l-S


U

c3 l-l


"o o


'5


3 O •-5


QJOO " CO
















13







r- 00







cSS°






So


c £

3. corn-^


C3

a


O


o



a


_


J


-C


f


H


'5.3



s


,0


U

U


^


h


'A


•3


b


o





c







c














b






b :


o



b '


c



















M






a






a




t




c



























3


X





























-d



a




~


ft






>



a







•o




C


1







"i



r

E


o M


3

be

03

a


c


5


3



2


3


=




c


u


a


■~


B


n





s



~r


-a


o


(M



« s




o


a


i


"JjocT





■*


a


■ Si 3


jC


T


§.i


ft



«« bD


o


<


-h oPQ'-'


3 O >"5


<


« n o




o


z





o





o


p



o

ft


c


a"


>>



o





3


.2


S


03




a







o







o







-° a




O.S




> 03




03 w







a -3




£>





o






CO



m— .



M


o o



o








coco



CO







a



03











03


>


§


a



CT





03



03


3



3



f.



<



_


Pm



S3



CO CO



CO CO






CD


Si


^J



3


3



Ph


Ph



^j


^J






« 


3



•""' CO


►-H CO


3


cd oo


gCO


PC


g>&


Mj









03 ►H


d HH


pq


o


o























CD >


>





•z


m


CO





■g


03



«-"


3



c




'm


GO









ft •*



- o


o






><


^ X




b




o






b


■b



o










CC


a o



CD

J3

O

3



C






S


.3.























ill




8



O



(■


/?



X!



c


o^.


h




3^


2&


fc


CO



CO



16


ETHEL BROWNE HARVEY










1




43


JO


J3


X!




a



3


3




Ch


(li


CU


Pn



w







o

K


c


cc £ co

CO (— ) CO


gco

1-1 d


Ceo



a



d d



d




.2 'Sc


'- — " "5b 1 — >


.2 'Eta



« 


©


H-l ©H- '


© .


cu 1- '




a


-T C -T


C CO





I*


CO £ CO


Eco


n<o




03


CO jjCO


oj


ojco




o


o


o


o



K







a


CO


CO



CO



>


p


p


o


p



« 







m


CO


CO


CO


CO



so

o


a


c5


fl


fi



©



o


CD



o


>




>




©


CD


CO


©










OQ


to


OQ


02





43


+,











to


g


a










e


_fi


©


8




-< S







a:


o


"o




"o


a


Oc





o


o




o







><


>H


>H




X


X


X


X




■b





<D


a


Ci






H


oc





'.2







g













C-!







o







ja


a

H

O


z


■fe

a






a







h







o


t


r o


b





©a



e










f<







DQ













6







z







~ 3. H





ex



O H S

j « 5





a






CO






o

CO









£







■^








si

s




OS







'5




>







m










03



~>>






CO

a 5

a


E o

CO

3


3 ©


a

©



tz "c




CO


03




« 



Ph








no


_o





13




"m


3


3



03




o


'S


'3






Ec

o s



.3 3


ffl




c*

3


_C ©


"3




<<


C



O



C



9



d


d


J3


3



2


•o


Ph


Ch



f


^J





o o

N


o o N


a

CD


p. 33

Inst, p. 33



§


js


«B - 2 B





3


V *




-*j



CO C CO




"* *S^ H

S


CO l*CO



n;


S3


O


oj

o



CM






OS


05


to


CO





o


o








M


M






a


X






2


a



13


13


o


©



a


c


>


>



©


©


CD


©



w


ffi


w


DQ





^


_^






™-o





B


H«S





B


CO--"





CD


©"B










O






c


°.2










>


>^X





X


XX


o






3






-r





'c


b




^"^SJ


CO









,0


2 1»





CM


08^





^

CJ






M



E




— CO o



a




no



mo




CM CO





4<-^<



CM CM CM CM




CM CM









E




C





M



CC


'c


ca



c


a



ci



a


"3



J=




a


a



t c


C3


T


S



'E c E



B


E



M


c


>>

)■ ■©



T


Si


c


^



<



£


C


)


CHROMOSOME NUMBERS IN THE METAZOA


17


■fi



a


3




Cm






O


£


co

M


N


CD

°M CD


a


ft


£


-o


O •-3


U






p



p


a



a


01




>



>


CD



CD





m



CO


o



3


"cd



d









•/-;


!*


03



X




13







.Q



3



b




o







'b




o







to



M


to



w


o



CO





03




a







Lh




s



03 bo




3


a



>


C3



03





"a



0)


T3













o



o


G







U



b




r £1


a



-Q




3 CL,

P-,








s *»




coo ^oo


N




.a a


a



MM boS


W





1-3



0300 goo



O U



CO CO





O



a a


c



CD OJ


OJ



> >


>



cd


^



.2. -*


—1



a a


13



















O



a a


a









>* !*


>H



X


XI


X!


g





■a

















0)













u









o





u





•"






b b


b



O O








b b


b



c


o>








SO M


bO



ft ft


a



cz


00







a







3







a




0;


.2































,a


3a



C





c









OJt* c




03


'3 e




a


o.a



<


H



O





CO


a 5

(D ~H

M ■


c« 


«3


a


CO





a


« a a" . s>

1>


^^


ft


00


a



^




CO


. ft r


CM ,0



CO

.a



a

uo"

OD



J3

03 "-5



-=


.2


c3



3



Nh


CD . 1— C


i-s


&


t-a



"a3



"3


O CD


"3

Q



"3



O

03



N


«i


IS


N


N



J








1^




oa




CO


O


t^


p


t^.



p

3



■e


p


03


p


c


_o





J3


>


.2


CB


u


C



s




"0



"O




'o3



a

3 O


a '0 >


Eh

03

a


3


02


a H


nj

_a



.a

CD

Q









-G


c




















O











•OJ


^










c"










"*











CD


3










"O.


2









X!


>

■3


> bb










•B S









£


X




|



"b



b



•b




i"



CO



01



OS














'0











/,





















t











3











si













b



b



■b









OS



O


























>r
























b



b









OS








"b



1-1



"






cr























s

bl)9 O ^



1

CO






bO O O


bt)

a


bO


CO


bd



bC






O ^


a


bD


a



O






"O


CO


a




O



CO

1


CN




00


m .—


00



O



UO




CO



CO



■*



CO



c



C3 ■

03 CD













3


















o5







co

■2;



.s



3






'3


"ol


i



3








S


d


c



_g



3


a





3


03


M

si



'bb



CD


c





'5


s


s



a



a>




co








.a


a



3


3


3


3



3



E >>




9


CD


CD




c












15










~c


>




>,


>>


>.



>1



O


C



Q



Q


O


Q



Q



J3


j£ a



3


3



F4


Ph








♦i







2oo

C co


Bg



h-l ^





ft ft






'&a


•flB ^



d Eco

03 co


Cam 36,

Jour 101













CD






d




OD CD



>


> >




CD CD






CO


CO CO







v-H



~H



c



c









CD









O



a



a












X!


X X


CD S




b


~





























w





-C




■b

06"



^



b





C3>








tc be



c-S


a a




























bll







a


c


3

CD




tH


C




a




C3



6







m



W


U



JOURNAL OF MORPHOLOGY. VOL. 28, NO. 1


18


ETHEL BROWNE HARVEY





o




ft


6


c.


o


d


ft


&


c



c


ft


j


















3






H


CO


ec



CO




»rT


»o



■*t





63 K

3


cs ft _©


09

C


d


"o

O


o esj


o N

ft


"E s N

c


CO

"5 c n:

o.


"r o


"3


N

ft


S

n:

c



6


1

s:


»^oo

ft



fc




CD


x





X



X



~z



X


© „



3


CM

o


N


H


K


W


s


H


"i


H


W


a


'E


H


oj 1 — '




O

OS


h <


o

<


c


3 O >-5


"" 5

>-5


3

o

1-5


i-s


'Z




oo ^



o


£ cool « 

c



^



pa








CM





CI










OS


CO



CO

p


~.


oo


oo



p


OS


CO



> PS


i


~


p


p


,c


O


a



p


p




p


p



K



ci








o'





b





n


>

a


C


TJ


c


i


,3


c


c


^


3


P


r^


c


CO

3


3




"o


o


o



01



IS


ej


CO



IS


o


CJ



c


c

Eh

U


s

c >



>


S


>


j:



>


>


>


3


>


>




Eh <


©


I

■7.




o


a

[I


0)


01


o

w


CO


CJ

X


© 02





co i









u










"3 "









o










£°




^


£



m



«2










-a c



C



a





w


a




c




















a;



•a-S



03


_


m



m



c




CO






3 E





CJ



£



.-_«>





cu




•<



03 g m o ^



"o


I


'q



"o



©"o


CD




"c




3




ft



ft



ft



"o c

"o




ft





■g" s



Q


ft





Q.O


ft



o













■w "rt 00 0Q CM







■ 3





>



>H


>H


>



Oj/2 c MW> *


c




>




H



X


X


X


X


X



XX XXX


X




X



a




































































a


















9 §

"3


g



b




b


b



b



£•*


b






< =


f


oo





•"'



c=



-2" 7


^ H






Q o.







rH






OS OS







O o












CS~







2 *




















,_,















w __.


g



"o









^b

Ss bbb

2 CKl M^J>







5 <


o


b

cs 1


2


b



« 


b



b


a


b



o


b


b



»ra


II







OS


00


OS OS OS OS




« 


r^


00





^- "


~








o




rt


~


HH


z



■b

00









oT





^





















g


i


~o


|















>>


CO


©


b


b


■b



b


o


bb bb


b




b


b



1


a


II


or


CO





OS


00


o ~- «n





£r


oo



91



■fe


b>

















CO
















6


















i

Q W O

S K P O H S J « S 0< < i


CC


s




bt


hO of



OX M it


to c




M




ft


O





ft ft



ftft ft w 03 ai





ft




CO


o





00


00



OSO —






CO



- £ ta




CO








»-HCM CSJ


c





0O



Q w O


















Q


















z


















^

































03














03





03







0]










9











©



T.


Z C


a



3

a


3 c3



1

3 03



a

>


3

,5


B

li


O

I 3 t.



« 




e


.2

"o


1

V



w

0.

a:


IE


3

|c


_3

IS


s

3


B

9


E


c



eg o

C3 03 . c .2


a e


C,



"3

S o


T3 o3





c

c

Be


- c c

1 > —



a

a


©

o

>>

Eh

3


C 3

C


ft.

o O


>

u

c

a


>>

CI

o

a. o O


c £ J

'Z


O C - u XlS. o3 a

So



c


C

a

c


a! "c g

§

c


p,

o Q


a

Eh

o

Q


CHROMOSOME NUMBERS IN THE METAZOA


19





i


»o



a i




XI


_d


o


=o


a



r Pi




3

















o





N


O


^



« S3




5f?


C co

KhOO


id


0,


a



ft ft




2^:


a


X


X


o



r*1 Sh-T




<D .


K


W


§



' '3)>— i

s




J CD




CD


CD



>



> >




>


>


<u


cd




a <d




CD


CD




5








02


M



02 03




02


03




.2 ft



^w


r



^



« 



O <U"3

a m a



J * II


~



«-a


J5t>


a


_g


03


cm


.a .«>



CD


aj3 '~"a


c2 ■- ft

co--« 


S


CD


jxjP

fl


cd o>.22


CO



jgHS


CDT3


c


"3


c -^


jj


"3 "gto

c, a_cu



C


o no


lo



>> X




o ft

03 2

- 3 a c3


a


■2°!

"3 00


3.2

><X


O S

>hX


x


2 "



V


X




XX


XX




b










cc











~~











<o




















b

£




b



b

b



^=>





OO




US


•*




2









"b



"b



£ ^




"b


■b







OO




lO


-*





















|


to a


61

&





M o ftm




a co


CO c


CD CM


co

CM





to'b




o<>


c§0 -00


CM


CO





rt




coo

CO


tl CI

a

CO OO






.«i





























1








CD


















c 1





cl

a








§

CD

~a

03

a

03 CD

03 -0 A


r




03


_6p




"3




"ol

60


u

C3


03

a


03

03 O



c

'E c C

B


o

O




'> S3


c









03


03



S




o:

c


C




-3


J=


"5


ft



o

13




"C



£



1


J




c





H


f




xi a


a


a




£ »;



eo








§

o


O



CcotS!


N


N




ft a



a







'3«« 


t


W



Cam

30,

Jour.

101



o














o




.


bO


.



- a


C


c



CD CD



CD



> >



>








X




m 02


02
















a






CM






a











o
















a













V




y.


o

B3





"b

o












K






a






a






n






i-l













o

QO CO


>b

o





o

CO

1


o








M





a



ao co






CD o r






cso'b






-<no>





ji


in






_3


.s >




c



-3


bo 5?




=



a

CD


c a




c

c



&


So.




s



3


CD







3 3




c >



a

F


.5.2







.= .a o



tt,



^J


a, a,



CD O S3

■ft'a-S


20


ETHEL BROWNE HARVEY


r

o

= x

w



JZ


a






£->


£


a



^


o




ss



H


Ceo *— '


K




a


2~ 8"


B


ge « &








3 CO ,_


U


<


K




a



o



o






a to

n

c


c

c










CO


£<



















PS


s



<




S


a



a


c



Q


"h


"fa


i


o


CO


a




f>




£




b


"h


■b



o


CO





o




z




cn




a




h




(o


"b


•b



c


CO





E









O




z




a g 3

« S H

9 b a


g. a


£ ^ z E ■*■ a a ~ o





N



rt


a




55




•<











a



c







o


B




a



"3






a


B









- >,






fc


C


)



one —

• — " a

" »£

& til





!


b


"b


b


c?


<*i






■b


■b


"b









H



a




X




«o


30



c*







3



3






■£


9





.—


c


o




>


3


o



-2


o


=


c




XI


>>


o


a

>


a

DQ


'_J


o


.=

=3


^ :'-'



C.-3


Sag


•S —


a



2 o a a


- OJ

— 9


o G.C

>H>-X


X



XXX


b g


UMao a c jo co


a a

co o


CHROMOSOME NUMBERS IN THE METAZOA


21


d -5



c





co





a


.






d





_T



























.



M















>->



N













o







CD











•fi



O





w



w



©



o



a



c



o



o



d


>>


.2 >>





o o




o


~Si












o c3



-3



"S"3




3


2 o.









~*


a









-i













>>





£





o





T)













o





£i






O



o









CM



o



O

CM









o„





>> u





tf- O CD .3








■*■*« 









ooo







£







c

























c5



o




H







O









d









§




§




■a

ft co .d

"o


N



Hasper, Ml


1

o

c o



o

d




o

00

1





co co

•S.2

a 9

O.g 1

O L.

3 3

££

° 2 a O O

1HL£



OB






A


Oi



K.





o





Ol


o



c>



"[





2 00


■■H


CO CO

•o





ft





d j. r


d


d a



i



o cm





o

o O


o cn


to" °


8 8"



='






ft

— r


m r 50



_ r



3





"= r^


9



"3 "3



H





H H


CQ



a « 



"o





O 3§


g


a ^


"o



a





£ 4


« 


.? C?


S 5







— o














2*



































flQ




CO CO



c





13



=


= 3



©





.

e>


© o




>

IS





> >

'-

> a>


£ >i


> >












oo





02 M


I


h3 H


M 00



J.g


S



CD


■ ~







cj » e3


c


o


o



^











s



n








=S g




o


B "" ~








b|



■=


~!


- a • •








C d




o .






^



""*


o


xj


II





X H



>H°



/■


^ O bl







H





IX


2





£ 55



b






b




b


b


cc






CO




CO


CO


'3





































r













U













T3














■b





b b



b b


b b



CO





to PC



r^


CO


r^


CO



■b





b b



b b


b b



CO





cc




oe


CO

c


CO










« 


a) MO £





M all






b£ M



Jl


MM MM



a 2

00 O






^. » O




- &=>g


ft c. O to O co



eo






CD CO



ME co "2 co


O CO CO CO











ft s

















■J





























c













CO



_Q













3















CD















"2



B













"S



3





£



r


CD



c



a





1 C



C


c













2


"c


cj



J2





IZ


'1



'i


"S,



g

s



"3





a



c


"ft


r



— ft

o





C C


- 1




CD



s



e






3



"5



J=



<





<


c




C


a



c


H


22


ETHEL BROWNE HARVEY


o


< 73

Q £

CM S

o .

M

<



a


6. c




«o


■o


»o



__;



_;


o



c







B


tSJ


s.


N!


OS





a


a


c


»,


X


ft


X


(5


W


W


H


55 !5


Pi

3


i5 bui







^


^


1-5


« 





>












a





(0 Q O


a


a


C


o


a



>


>


>



o


c








w


ttl


m



^


^


w






93


C


c


e





X





K




as


■<!

S a

K








C.


ft ft


o



o






><


>


>>



y


y


X


Q


■b



"b


1


CO



CO


a





H





ft<





b


"b


V


"b



to



CO


Q





z





CM





a









■b


"r


"0



CO


CO


CO






« 









o





z





fiSn


Sr 60



St to


£: ■< t\


ao



-2




co O


CM<N




Q o


" ' "


«-*


a





z





-<:





















o






+;



o3


a


a




£3



1

Ir,


a





0.


03


c


s

o



O



-c



ft



CJ



■~


c


J


C



ft


a


a


a



ft


c


c





»o






r^
















o o N











o




•■


o




c



o


o


o



o




c


S!


tS!


N


CSS


rs.


N


s.


SJ


S!


ft


a


a


A


A


Q


^



c






■/.


y.





y.



m


P^ O


W


W


H


w


S


■~





3 O




^


3


■o b us



- S





1-9


i-s


1-5


1-5


n


>S


-;


>-!













o










■>J<




■*




-*





r


a


"


~








.2


>


-2


J3


-2


2


-2


■2


K


N




CJ


o


U


a


CD





S


m


S


<3


S


~


S


~


S


f'.


-CT3


_ o

-s is


T3


Pi K


T3



















o


S°-~


■■



c



c












ft ft



a



a

















-o-d


CD 0J



?•"■ II








'bo


a


o


^ II Jl

a — .


o


»


t>


o


o


o


a a


a


L +j ~

c


^


C3


r



a


P*X!


^ s l


y


= a o .a - "

a.


X


x


y,


y


y


y


yy


yy


y


X


V


y


y


y


y



b










-CH










b


b









' b


■*?•


•*









>n



b











^*













M











o








so bo



SI





St


ft c


a o__


o



o


c


o



c


c








5





COCO


00 00 00


00


o o


oo



00


OC


CN


o




c3


03







•a


















.a


-a







c3




o3

a o a


a _o

a a


a

JO

a g


"fi

o

a


03

03

a '3


ct

z \


i

3


c^


< ft


O



03


03






o


« 



03


03


03


03


03


T


03


DC


^














.a


.a


j=


_e


r


J=



-a


-a


ft


a



a


a



ft


C


a


a


O


o


o


o


o


C


O


C



o


o




o


o





eS

















_

















CHROMOSOME NUMBERS IN THE METAZOA


23


c a





lO







o


N


SI


c a


>



d


W


>

OS .*CS




i

>-5




c


o


c


a



a>


S


>



o




a


03






p


a








c



ft ft



o




>


>H


X


X


b b


CC


CO


b b


CC


CD


b b


CC



bfl bfl tr -C


CiP










T


5






C


03


01


03



1 S






C

c


' g






a.


h



MOO

ft en cc



-1




a


H


c

i.


£



r.



V



pq


03


fc



^j




^


J3


a


fc








0.



fc


ft£ ft


<


©~<U



OS. CM





° a °

CO gco


t~


H




c


p


!* £





CD



£


c



bfl




c


a


c


o


%


S


X



c



<M



c







c



a






>



y



b.




b




b




bfl


ft






O





a>



a





a



3'


<H £













J3 C





'3 « 



N

































J3





X!






fc


~



fc



fc



fc


fc



fc




u


33


Si 2


"5 s


03


os



C


^


•o



ti a


e


&


Sc


Is


B


a


gca


c.


gft



<o


<


rt


<U


-i



<r>


<


^;


<o



<M



.


CM


«! 03



7 !


C^



?i




a*


a



2 §


cjlS


3


c




o


a §



a*


$w


gco


H^


a


M


gCQ


e


n


gw



H


H



fc


fc


H



H


fc



fc • .




-^
















o


^



o


o



c



>>


>>



>>


>>


>



>.


>>



^


















OS


09


a>








a


fc


P


s


B



£


R



E




o



o


o












M


bfl


hi!




















c


a



a


c


C








o


o



o


o









g


§



« 


s


§



S


g



<-.







yr~


o



-3










3^

— ' O

IT3



O










XI




K" 1









X




X










"h













QC




"h





9j













tS

























a













03













u













si


b

CO





b

oo




b






b




b


b,




b




b



CD





OS




oo






















bfl













ft









bfl













ft





CO









o

















O ft


















§
























03






J3




3












ft



















3




x>


ai




3




C







"3




bfl




"


ft




•c





3




3



M




03

a


03




ft




ft






ft




03 o




03 o















5




i!


3







^






a













a>


>>









c_





J






fc




fc


24


ETHEL BROWNE HARVEY




Phil.

137 Phil.


5 2


ft


3 5


6, p.

. 514 uppl.


Phil.

137 Phil.


CM CO


d


~


Oi




ph Pw


of


d, Ph


^^ o


CO


d


03

ft c

- <



H o

H

«  a h

m

K


mer. p. 154 6, p. mer. p. 97


mer. p. 154 mer. p. 97


"o o

si

mer. p. 154 mer. p. 97


Zool.,

, 14. p h., 15 s


mer. p. 154 .6. p. oier. p. 97


O>o

o

c

• 03


CO

6

a




Trans. A

Soc, 20,

Biol. Bull.,

Trans. A

Soc, 21,


Trans. A Soc, 20,

Trans. A Soc, 21,


dft

is


Trans. A Soc, 20,

Trans. A Soc, 21,


Jour. Exp. 69

Anat. Anz.

Jour. Morp

p. 223


Trans. A Soc, 20,

Biol. Bull.,

Trans. A]

Soc, 21, j


Jour. Exp pp. 371 p. 1

Science, 25 191

Jour. Morp


Biol. Bull.,

Amer. Jou p. 279




—* -# CO


— o



T-* CO




—> ^o





2 2




p pp


p p



P P




P SP?







K







coo


" »


o




t» w



8


>, £>>


>> >>



>. >>









>



u u



~ u











lO


CD CD


Ci


CD CD CD


US




-. TJ



K

a


s as


S E


p


a s


P £ £


a Ea


p


o


z -1


SoSo

03- &



o o o


o o



o o




o o o






n


£0 bO bO


bO bO


c


bO bO


g


bO bl bO


C


d


a




o


j -^ -^


o


+j -^



o


o


o




a co


CI c



d c



i 3 3 • 03 c3

PwPh

C C C








o o o


o o



o o



o o o







§ ss


s. §


fe


a §


%


a ss


£



&





■gg



■ss



«.S &


-o.S





3




XS



T3


d-s



S CD C

2oS


d

CM C


-73




.2




a


d


CD



o


C






to


CN


c 2



d K



-c ftg 1


d-^3


CM




>>



« 

S

w

K


c


.3 t. o


_a


o



°o^


Is

ft o


_d




o m



©

ft


CD CD

"3

ft CCD

Ho


CD


,2 u

"o

a E§


n oog

small

ipg; t

and


"o ft




8 J

CJ O

O CD 03 "o


-3

CD

3


p


o o

o


5 2"


XX i

Pair in i 2nd ate


s a--©

op


o




3 O (3



X


M


X


X


S*i




Z


a

a o O


o B


b


b


b




b


b


b




b


1 Q « 


CO


CO





o"


o


o*
















o S














a, u














o .

tf CO


























n


H













H














tf


o


5 •


5


b




b


b


b




b


■<





t^




s


s


s




^


,_•


a














CM














w














E*














O


5


b


b


b



b


b


b




b



'



i^


r-~


S



2


J3


S




m



t













03



























6














is














2 S g


bO


bo


bC bfi


M


bo bD


bd bO


bC tr




bO



ft


ft


en O


ft


C

c


ft



&8




ft

CO



°< < £



CO


CO **



c




^HCM




CM







CM



CM


CM CM


CM CM




CM



£














<














DO








C














o a ft.

CO


§

c

3


"3

op

'ft


"3


CD

bo

E



b

]


.2





.2




K




o



b


t

"C


'E





"C







03


C3


+j


-m








3


3


3


03


d


C3


03





03




-a >>


>>


T3 >>


03

d


3 C


03 C


a

c


03

d




I




5



<



<



<:



<!


<



<



<




<


1


CHROMOSOME NUMBERS IN THE METAZOA


25


t>r
















a


ft



2,





CM





ft


d




J


OS


CO



J3 Ph


re


la




o>


a

CM"


to


CO


J3 ft


-3

ft,



r





CO


P3




CO.

a


ft


05


d


•*






d



_J


CO



T*



§S





_ r


CO


d



<


oT


>TO


I^r


— —



fe


99



OS


"o


o

o


6


h


rH Pi OS


N 10





o o


6

3


oT




rt _


3 0.


CN


fi^


a


p,



-*


No


N


a


5


&a


ft


-a ■ c





N


rt



3 o


-i


pj:



^o



<!




~ r


d-o


d


CM


<


=:<


rt

ft C3





ft


lO


— '




"3



3



a





3


x a





r^


?4


Wrt







3



m


to r

C3"


w



pq





« 


w §


H


g



r •


J


"co





CD


m




"o


«1

osPQ


~c



"o


a


3



"o


1-5


^3


fi-H _CUOS

CO


3

2



o

3


s' d

3 ft O 1-5





i-a


a« . •22"o



<


S


V


£


H


BH




S


H


H






CO


o




>



.a




•**CO








rt


CO











§



s



p


op








o


p













TJ























•a



a



>>


>>>>








>>


>.











a

OjOO

o


o


o3

m2


■^


CD

a


Pi ~

fl> IP

SB




o


»o p


O


p


a


3



p





Oi

p


p


o



a>"



b


o


o


c








o














3 .


So


M bO





c


a


a


a


SO







a


a




> —



R >>



J

-M






o


o


o





O





o


o




-la i-i



O o


>,


c



a








3


3









p.



o


o


o


o


O





o





O


O









o



§


§


w


a


SS




a


N


N


£


a


§



£





&


£


s



-a


■a 2


•a



h






•— _o_o

■g'fi.'Q,





■a

3




_^ o) i

CM 5 >>T3


-3



a

M M



"SJ2 33



CM


•5.3


cm



o






^•3 -3


"




CM




a s

•" a


"


CD




'-jTS




0."°

^3o


a



°o






a


c




3





CD





o

^>r3



CD


<a



E-°






CD u —


a



M


S




'o co

ft. 2


>


X g


Pi CD


bi


M



"o


"o



o a






S H o M


~5




3




if


1


1


ft



a



c o


b n ■9M


a o








ft o




ft O




si

co


a --3 Pj CD 03


a y




ax

X



X


XX


M



X~






X


X




X




X





M



X



lb










b


■b




■b





















00 O

-00


00




cc





















t^


1-."















o














<M













































b

b
















o


b








00 &


00




■b





















.00





ro





















t^


































































CO
























o


sl



^






'bo


■b




b

re





















oooo


00

















O
























_

























"o u
























bO


b<>~

<M CNJ c ^


MM)


s

o


t/ bfl






M b0"o-3


60 bD



hi





bO





bO bO


u


bo tr'3'c


CM CM

bO

ft

CO


a


c-9



°-9






»9^


GO



ft





ft





a 9


c


S-Sk.


CO



CO O


CI


CO O






en O>go


CO O











co O


GO


co o bo



si m °



CM


H CN





IOCDU3CO


lO CO



»o









W3CD



i-iCM ^HCM


CD

a o


• CM


P, O CD m O B

-NO


CN CM


ci


CM CM









CM





CM





CM CM



CM CM CN CM



cmcn -—









































c—






















1








pi








pi













H








^





O




O





o













o






B








03




03





03
























3




a





a


03


cj











B a






"3


Pi



03 Pi

03

o





3 CD ■p»

3




a a





a a


Pi

CD


Pi

CD



^0





a





^







o

2


"3





03




03





C3


>


>




1


a


|


d






3


3

Pi



c8

J3


3 Pi CD




3

Pi CD





3

Pi


03

CD


C3 CD



n

'pi


'C







0i



CD




J








~


T3






•tJ


c3




|



a



p













03


03


03


co.


«'fl





o



C


3




S





.2


'S


'a



id


§


03


S


ro p,

03 o





IS


II


2



O


C3




03





03


o

"3



C


e


B


c


fl>a





H



Pi



ts


A




A





J3



J3



<



<



<;



-1"



-t:






^




<



O



O




o





o


o


o



26


ETHEL BROWNE HARVEY



■ r


^




a


& a


a •- d

a


a a


a a


i

a a



CO t.j£


IE _a Ph Ph


to


CO


S ^„B^


CO"


r* to

CO


CO


a


— . jo CO





to »


■* x"



=o r


o r

■m"


to r


S3






- 10 • a -ft

^ "B n t-°






O

55

a

a


■38^55


o . o .

- - E

o o


2 I <m a


o O


8 1

S a


a N cm a


-a

a


cm a


fc,



CM cm


i7


X


• r .^CQ • r a " "1c • a °

gOC! £Ph*2 u.'- r H P- P3H



X


X



x


a


? a, c r- x


StiSc gogo

- CO g OQ

- H



g H Sobo 02 ^



-- W


Is^ o o

CO >-s 1-5


- C-h .-tc

Jffi _

CO ^



— "** to

p pp


p o




p p pp







K



~„ ~ „




..






a


r*> >• x


>> t»




>. >>>>>.







>


1 si


o c


O


r~ en


O V 4» ©


CO


r^ c^


r^ OB


pH


r^ c.


a


a a


p


p p


= S aa


p


o o


= o



o o


01

n


C O 00 to tl


5 5


a"


a" ■ a"


U tC U M


(3


C


a a


g


a c


c




o


o o



5


5 c


o o



o o


"5 a a


"a a




"a a e a








o o


o o




o o o o








% gg


~ ~


sS


% £


g S "r-!


==


>; >:


& 5:


=


£ £



_; a



a


a




a




a



5 o



•"■


tm




•^




"*



• r T


~


X


X


-T3


-B

c


X


a



X


10


a g


cm


X


X


« 


c*


X


CM



X



r



t






S



©




eo







a












H ><












b


b




b








r

>



t






Q












Z












CM












a


"S





"a







E

a





a







1



b




°0 °




b




F

- e9





-^ s







tr.


b^











c























tc



it to


M il


bC


to tt


to to


to to



to to




a c


~2


a


°-2


a c


a 2



a 2




71 O


in O


m


a O


en


co O



00 O


< •«! £

cc



wo


HN


CO


co-<*<


CO ■<?

-^Csl



COtje


S 2 w




CM CM


(M CM





(M CN



CM CM


2












«!






















3











00



a


a




m

a




r



S


U

o


-5 13

CO


_a "5

X

o




a

3

o



G i





03


en



a



to


o a

a. n


c

CD


g


"EJ

o


£ to

2


CD

00 CD


01 0>


'C


a

n 3



a

8

c








n

o





1



o a >> u o

o


o


CO

O

i


5

a


o o


1



00

3 M



c C



c

t.


)


3


a

w


a



a h5



03


CHROMOSOME NUMBERS IN THE METAZOA 27


CO ft



ft












ft














_ a





O co co



d












CO












J3


Ph


CO

coco*





dj \



_J












o^r












TH



ft— r





m o



c












o












JIC



. o





cm O


























fe


CM P





— N



SI












— N












Id


S a


— tSJ





S3 d



ft












— ft












^d


<~


d c.





3 *

raw



a






















CM

c? si

5 p gco

H


CM


3 x





O 3 S3


-r


1-5












o 3 2












go gco

H


O 3


■>)•



























rt


CD






























o


O




























































h


>;






r-~Ci



O












t^o


















op



r" 1












PP












E


s


op





a a*



g












a a












o

60


o

60


a* a





o o



o












o o














p p





jnjfi














CO CO












a


a


































































%%



£


-a a

a

V

3 ft o

-M











fefe












S


a


5S0S





-6 £ a*3

a

o


CO

.5

v.'

CO

a

c3


5 > o


a

CD

3

a o



m

13

a

OS


yj 3

03


S co


CO

— a


O

s


CO

CO

•a a

ccj


9 O

3


CO

a

CO

■<*■

■a a

c3


O


a

CO

.3

CO ■0

a


DQ

2

CS

Eh

O

5

_o


St

o

"o

ftp

p


CO

p


o

"3

c p


a*

"5

a o


a

c3 co w 3

CO « 

c "gS

C3>2


a

^0 ^ 3

co t; «3

tj£ a o



J:


>H


>H


a


eg


!*JL


ftK.


a


g


09


&M


><


s



s


>^S


>hS


"^fe


.5


>*


>^


>Hg


X




X


X





X



« 


X



X


XI



« 




X



X



X



X A


X


X


X


b









b









r n














b


eq




b


« 





C4



b




b


b



CO





b





b



b


b


b


CO CM


"^




— •







J,




"- 1


^J



• H





i





d



o


-•


^


"


o





























































•b




b


b

a i





b



b




b


b



b



b



b



b



b



b


b


b

CI


b


























































































































-a































.2





























































-is




b


b





b



b




b


b



b



b



b



b



b



b


b


b


b





cm








CO




CM




-j


'


CO








CM


CO


•*J<


b





























































~*































60




60 60


60 60




60 60


to tc


60



60 60


60 60


bO bO


bO bO


60 60




bO



60


60 60


bO bO


60 60


ft




°-s


ftQ




no


ftO


o o



0,0


ftQ


ftP


ftp


ftp




ft




O-P


ftg


0,0


to




X o



o





o


co O



to O


CO


Q



o


CO


o



o







co O


co O


CO O


CM




CM CM


CO CO





■*


COCO




CM CM



S


-r


LOW



CD







CM CM


coco


■*■*


CM




CM CM



<M







CM



CM CM


CM CM


CI



CM


CM CM






CM


CM CM


CM CM


CM CM









CO

3












3 O







.2





.2 :










-*J

















"c5





"3










£












3







a





a










o

S












a E




















o)












60







CD





09





















O







OB





00










|












9







3





g








































-3












"3







•3





'-3



28


ETHEL BROWNE HARVEY




0.


ft


ft


,t


_J


-h'


ft


r~



Zool. ISO

p. 439 bt. f.


ft a o

CD ^h




EM


— to CO


d


2 s£ ft, 2^


CN ft ^


_ T.

a a a

S*

a M



CC

6

a

era

a

o


! CM 0.

H


CM* CM

o


■* .

3 .„■« . S a °.Eft

CM 3 CM


r cm" ft "3 w -r

ftftg 1 E3

x . o a


«> -jB'tseo


tS s

ft 3= X 3



0)05 JI/C35 go> 2

M IB h, ►,


gas o gco

h en


S 3 pq


. S ■ • ° fe o o^


^S'o







.-t -^ CO










p po


co




a







p




a






^ >;>;





>



r~


r- o>


,_<


Lh u i~


■a th o



m Oi


a a



o


p p


it*


S ES


p .-> ;*


■^ -^ CM


p p


CO






o o o





03



B


C 13


c


&o mm




B C


o



c


O O


o


C CO

o o o


8 ° ' E


o o o


O O







[3. 73. o






5


'=£ £


5


a aa


£ £ a


o c5o


£ S


a « 


.3 .2

5S en (Q co

■5 9 ^a

t/2 t-. rrt (-* MO ^O

T3 1 T3 §



X X



■3

a

CM

a


■a ll


ag 3

2 ft cVD o

X a

^-jo

»«£ g


•- s«s

« 3-- 03 O." 3 ft O || v$


•<

a

a



bt a



o

"o ft o


1 \°


XSE

« 


g.2 C <2

03 . ct) .

X X



a m

.« o

o




3B2.C OCM CS




X



X


X X


a


CI


Q


b





b


"b d


b


b



lO





t—


t> o



C<1


s


1





co"


co a


o


o"


a








X « —

r ^— CB


X.S^j


H

u

1


b b 2





b


b


-1.3.3

O o, II fto?


ss'jsl


CJ









~~" V


z









btcjS

o


a










H










X










o


b b



b



b


b Oc

b


b



U5 CO



00



t

t- r~













H










10



















6










%








c



a a o hHS gfca


MM MM


t£ M M



M


MM MM"5"o


m!


M M


n.o ao


a CO



ft


ft S ft ° r


ft


a 2


CO O CD O Ifl to CO CO


00


01 o IQ CO



CO CO


CO O CQ O^O


coc>


CO O CN-«1<


CM CM CM CM



""



~^


Hrt "^ ^^ ^^ ^^


CM CM


CM CM


K










<











' ' '












CO













3


3
















a











cS


03



c











C


c












oo


1






o


CD ©


M


M


a






bo


M M


h


h


g






03


03 03


03


03


a


CD



oo 3

M



g


1- c


E


£


p<





"3


"3 "3




on


3



'5



-Q


Si JD


CD


cu



O





O


o o


1


a



a


1



a o


a a


s


s









O




01

a


r V


c




c

a



c







m

c

03

Q <

c

P4


s

CM

ft CO

_a

IS



p

>, u

cu

a

o

M

d

o

a




s

o O






b

CM





c.

■ «  i '1

c

C



CHROMOSOME NUMBERS IN THE METAZOA


29


-3® a-*


X


3-S bl CD

"30

a



OB


CO

coo




— '



— |


~



COCN



ci


IS


2



IE


13



CO



<M



Ph


Ph



PL,


PL,




&



d









a



P.




«y




■*(<




o



o



'C




^uo ^t


CO



o ■c*



S a


5


d


Oj" CD 05

S dS a



3


3


si 3


CD


s

<!


_r


^s



'3


tS


<


s;










4


_;


-=


5. o


3


o

Q


1° So



3 a>

l-S


a



o



gcQ


2 w



<


N<J


H


H



H


H








CO



rt


CO







o


o



p


p







"



"










>>


>,



^


^







t4


^



h








0J




CD




O


cc


ZJ


"


a


E



|


S







o


o



3


o



CD


CD


cd


"cD


bO


SI



bl


bo



MM


_«.*


a


a



a


d







o




o


o



'&&


%&


a


a



a


a



'o



o



■y







s


ci




_ <D




a




o


a























CD


GO


Q CJ







o


CM



"3








II


'>


a S


cc




a




bO Fh 03



-3


>

-2


32


a



o




J



><!



><





X



a>












5s






b






'E























CD

£






o






o












c












■3

































CD













b



b



b




b




<M

























-3












'i


-f









b



&


"3


5




s






eg












£


03











bO












aci









wi





em







0.





a










-Q











i











c


D,


c\
















§















.2^



s







03

a




"C



s —







'3>




§





c




CS




03



3







£





p,



a








03



03



2





3




+J



-*>



_E








cd



0)










s



£



a





M




o



o







O




u



H



e





3




■3



-3



Si





£




>>



>.



>







« 



=



M





3













a a


a a



13



3



CO







a



Ch



Ch



-M CO


Pv



Ph


■^


CO


CO CM




■^r




03CN









o

a


d S


3>

d


^d

dco 03 "5


Ch

CS

E


a


CD .

S ft


z I

csa


8


3fe3



<


c;'


-1


M


«!


p"


03


a a


ill






r




.

W


H



a H


5


a


O



cj-3

gdH



ci

1



c o

l-S


c» h-*

CO gco

>-5


1-5 "-3




CO



^




CO






o



a



o


=



o






>,



>"



>>


>>


>.






u





b









CD



03



CD


c



CD


33


CM


»o CO





S



s


£



&


p



O p



o





O


o



o






bl



bj



bO


bD


bC



3


3 C











c


O


o o



a



P



fl


"S



a






o





O


c



o






a



a



a


a



a


5


£


% %




2

O





a


c



■a c



-a a

CN]



CN


a





a


^






.s






_t




c



.3



_o


o

o





I


c S



a



CO



"3






"e








o


E


CO





CO

o


a o


o

a




c






c





o




C





a


>*



45



X






X




y



^ X


o

o3







t




>













J













t>^














'b






«o



^



■b



■*






oo






























c













.o



















■b


t




■fe









ws





00



O c 31




















bH



bO bO


M bl








a



ao


ao












o


co O








-.£




■*■>*<


•*•*















1?










'c



e




§





■5





c



P



bl






f






3



03





_2


£




IE





§





3


CD

— C






3 CD



s





S




3

t



OS

bl



>>





>.





>



>.



o





o








uJ



30


ETHEL BROWNE HARVEY







a « 


— ' —









'-G t-J3


-C -C



.£5 A




Ph «pu,


Ph Ph


CO


Ph Ph



a u



- ca .


o o O N


cp o 5 ""



a

K H


a o." s &


E aSft


N


s as a



^"^


^sfs


a x


<o«-



N3 CN



£ «58


CN <M



2 o . e °

gM.o gx


1 § 1 y

gco gM


goga

•-3 '~ 5


2 cJ 2 ci

S o S o

gM g0Q




H «H


H H


H H




-H Hj<CO


— CO



^H CO




O OO


O O



O O



« 







a


>> >>>>


>1 >>



>> >.



>

PS


- " t

-


- t4



0) (DO)


CB CB



O CU



ass


s s


P r"


a a



QQ


o o o


O O



o o



a


M il :l


OD bfl


c a


it tti



o




O O




c a c


a a



a c




o o o


o o



o o




s ss


~ s


£ j£


a a




c

Ol

_c e

ft o


B

CM


•a .s

•S § f


•38



K

S H


"o a



KY to poles (Sometim and Y e size)


o

>* S 2



3



X


X


M


a


















a







o







O 1


a


b





<


i


eo


b


b



a




cc


OO



o







5







o







ti







M


H






EH


8h 8" u







b


b


b


b



t-~


00


00


t^


,_;


a







<M








— -u




— . 03



a


"d *




■32



&

o 1

Eh


1 •




a«u



.■3 1-5

-c 2


b

o


b


to 03 ca



1-1





00



6







z







H O


tJ!






"BE

S 5 w £ P« h; <^


a


M


M M


bC




a


ao


a



CO



CQ O

CO co


■c*



H c W

a * o



"*

""*



a







fc







.<












03






3


3


cS










en



2



>



W


tij


'3


'3


C



o a Pj o


O

C3


•2


03 3


03

C3




H












o


3


CB


CU




a


a


a


a




O


o


o





T3 ai


a


a


"oJ




C



O


o


a,











A




CL,


-Ph




03^



6a


Acad, a., 53, . Am , 21, p



<N



§§




gco


oPh gco



h


Ph H




1-H CO



o


o o






>>


>. >>






S




s


a a




o o



M


faO M






CI


a a



o


o o



s


a S




13




S




C




"H




4)







O




a










5>H




« 


03



b


-o







43




rt




55




00



8cf (= 9, X and Y at poles in metaphase)



b


b



o


o













3



5


03




3



Ci C



3


03



O


DQ



C


43



c



^




gcQ





■*




a> —






00



ft a 8




otSJ



CS


CM



o _ : a






5 3 H






c


_• c .




£2~



>>






« 




c







a.







c




a


CB CB




a a











P.


PQPQ




TJ




c




CM




a




•"•




CB










a










!x




X


1



b








CB




a













55 o


c


sl5




<n CB





W


n oh/^






b . u^5 C



o


CM




rt



b



c


b










C




c 1


J1




3






b


.S



a


C3












c




c







e


e



V


•z



CHROMOSOME NUMBERS IN THE METAZOA


31


-*


a




OS


»-H ^


^H •




ftO

o

ON


og


Cd


M tS


=" p,


d a


3 H


3 X

PQH


-h u .


,-J tH __■


o 3 a


O 3S


£3


>[edit] © co


O CO


r"T*


r^r^




cd cc


a> a>


a a


a a


•s &


if e=


o o


o o




PQCQ


mm


-o


-s



e



cm


d


c

■■*


.—


a>


3




o


o


a


a


o


o




tH


>


X


X


b


b


CM


CO




b


b


<M


CO




b


b



-■*




M


u


a


a



CO



co


<n


CI




ci"


ri


a3 13


u


d



3



cs








c


a



o




o


o


2



fc


i


a





_




a





a












CO


CO


IS




13


£


a






13




csi


©


CO


Ph


Ph


c-r


Ph


Oh


CN


CC


Ph


Ph


Ph


Ph


■*



r


a


a


■*



_r


."*




■*



-*


^


a



o

tS)




fc« ~h cd .

£ ft


E a


z


Sa


ad


"o


O

o


IH ^H

S ft


£d


US' o o .

3 a£ ft


o



a


_;


-J


<<£


<~


a


o


<~


a-a


a


<o


<_

<o


<« 


^



x


3

~%



CM


X


_Cfl



x 2


y.


Ol







w (a


m

"3

s


co *r

go

gX



s

o



2 ci

5 °

H


K

o" i-a



"H


gtM


S3







CO



_


o





CO



CO


2






p


p



o


p




p


o


o


o


















M






















>>


>,



>,


t^




>,


>>


>>


>•.
















h



dCM



>ra







o


c>


»c


co


o


o


CB




p


p


p


g


a


p


£


g


p


p




s


g






S


o



o


s




3


o


o


o




c


c


c


M


M


=


bH


bO




bfl


tr


bt


bO




o


o


o




o




o


O











d





c





C


a


c


82

&H






o


O



c


o




C


o


o


o



IS


£


&=


s


s


>


s


s


^


^


S


s


«5


s



-3 G


lis


d

CM


■Hi




1.J

« -g



■a



— d 2 o





-s




fl d CD


_c


-. o



d


C CB





s *







.S'-H L*





,pH


'"* u



"*



•-' ^



.—



,PH



J8


p o o


« 


<n O



B


JS



c



co O

cu c



V



CD



"5

a


ft a .


"o


oE



n


o




a



"3 a —



"5




a



o


O O Co


c


CO

p


O


°g


CO

p


c



°i


CO

o


o



o




-*+* o


-S c




J c




w o







>-i


r"Xfc.£


!*£


,e


>i


^-l&H


.2


>



3


>



>>



X


X


X!


X



y,


X



y.



y



y.



X


03 T3


b


b


b


b



b


b



b



b



b



b


■§


CO



CO




t



r


oc



CO



CO

















_D



CO















rt

















+j

















C CD

Ph
































,_;

















~


'b


b


b


b



b


b



b



b



b



b



CO



CO





t^





s



CO



CO




CO
























c

















o









































b


b


b


b



b


b



c: _



b



b



b



■*


•o


©


00



CO


CO


















3_>.
















































b"*
















00









tr: tr


eO cX bt



M bC



bC bC


M





t[ bO



ao

CQ O


ag a

co v.



a a




a







CO CO


CO CO -*



-^*


-J"



H* -f


CO





C<» CM




M

















"














H



3


















































c

D


'c




c








CI



C

CD



3 S




od


T


CO

C3



a


J



3



"



"rt



e-. O





|


C



z








Ih






"3

03




a


2





a



CD



3




c

CC


£



>


■ i

c



"3 3



4>

a c

1



e3 CJJ






C

a!


C

CC


O



~2


CO

o



CD

O



o



3



§




a



CC


m





c




O




U



H



fe



32


ETHEL BROWNE HARVEY



<N



^


d


00


d d



^ _j _: 55


d


io


d


—•



—•


d



is"

~3 O




■^


IS IS



2^13 IS pt^



oo



Edac


IS -a


a a


co

o N


co

"o

a N


a


Ph Ph

£d£ a


CO

"3

o


. or* ^ ^ a a - "* d~

^Cl . »0 . t^ m '£h

=> o . o^m

"o o (S3


d


CO

"o

O


Ph Ph

E aE a


P3.

a


C, & ■ X _, ft

. d u


d


d



<„-<_<

d


d



d


<o^-h

a


v.




cm c<)


X


X


"3



Cq (M


C»l CM



W


H h


tS! rd


03 o " C3

5o»o


W

H


^03 "a " S

_;_: a § a §d „



-a



03 t; 03 r\

Sogo


® ei m ej

2 8 2 °



ft §9 so

,0 1-5


3.-,


3


,_, o


gco gco


3rH


O O cirri cjr/5 o 3 NNE-h Eh -<>-s


3^ o

1-5


&


O l-S


gco go5


£w gM



o

>-5


O


<


H Eh


■5


Eh H


Eh H






O


•-H o



t^O0-H to 0»-H



o



-H CD


— (O






£


o p



OJOSO p ^H^H





O O


p p


G3





05






M





a






>> >>



>>>->> >. >>>,





>» >>


>. >>






u u



u U U U U (h





U. t-r


tH U


>


lO o


O


cc


"0


QJ 0)



O Q> O 01 O <D



"0_« 



d> a


o <u


K

a

n o


p p


O


a


c



o


E Sa S S3


O


03

p


E E


S E






o



O O o o




o o


o o


c a


a


d



_- bj bO



M bJ) oJ) SD bX bl)


a



a


M tC


bC bl


o p


o


c


~o o


"3 a a


o


a a a "a as


p


8^


o


a a


d a






-Q o o



o o o o o o




o o


o o



L£ &


%


'i


a a


fe


aaa a aa


ke


£


a a


a a



-3


c



T3


■n


_a


a .2 c £ N o So

m o o x

^§£« 

o Ph a


T3


-3


d —



d



c

CM


X


a

CM


d CM


a


X


a

C<1


a



a


on M K «!

S a

B


a


si

a


X

c


"o


a

CD

"o


X


03

"3


a


.2 fl

03 in

"O-th


.2 2

03 O

03 O

"5


o

C3 tH

o o


o


a


B

c


o o


a o


a


a • o oSga


c o


a o


»a O C3 g


ft CCO

o go


>


>h8


>


HJ

tH


>


■*H 60

^8



>H


>


^Xg 1



op



X


X


><!


« 


X,


X


txj


X


X


X


X


£



b



■b


1)


"b



■b


"b


■b


b




i




t~


t>



r^




l^




a





















9^







£




"b


b



CO



b



b







r-

t>



'b



t ^



'^



O 65














CM














a








_







Eh I" O








<-H







b




t


"b



DO



b



b



'


oc




X


■x.



b



oo



00



EH DO








3C




















6








_







•z








S







»-. ffi u E; g J G3 5T

Oh <! m


M t[


if if


ati £



m bt


CO


bC b£


E



bl


bt


£8


£8


C. c 03 O X



03 O



03 O




a

03


m


HHH "<*•


•*•*


-?•& *t



h* -^


tl


hh<-h<


H#



■«J<



S*o


























a








■z







<;






















c- ■
















03 ■















s


3


3 ;


03 3


3







00

a


a_



§

.2 'C


>


V

' z


•1




]o


3


a





5

a


tn


o


c


9


'C


n

03

>


c3

>


"3

o


Sp

_3


c3


&





tr


03



03 a


03



m O




CO


3


3



3


3


a o


3


3


^3


03





g



a



.2


»&

.3


.2



•a


ei



G

-I



o

3

H


c


IE

1


a


IS

3


3^ 63


IS 1


3

a


3

a

a


u

c



a 1



CHROMOSOME NUMBERS IN THE METAZOA


33


— : ^ ~


o". f. £ 2


W H § K S K


SS 3_ 3


S a£ a S =•= o. co


-"o^-r <©<- S


>-« 


w cq --5


m o s o

c o - o


g X 7- X Eh Eh


age.


£ tab


t~- ts>


. - .


. c —



d o .




gO a


goa


gee gec


Ph


P4


H H


W K


£ & £ £


a a s a


■da

ii


c-c



_oj S


X


_; c £.2




-2 =3


-~ W jg



5.2 o






5S


c3 O



S" 3

n °



3 rt 3


J2 o


T3T3 O O

CL,Ch


> 3 ™!2


3 >


JOURNAL OF MORPHOLOGY, VOL. 28, NO. 1


34


ETHEL BROWNE HARVEY



&i n



— ' --;


a



c



IE f-lc




a~*


ft Ph £ft


of


H o

a


•WO ■


e>


.ira a .J;

2 •«> 5J ■•

a a a a


"o o tS3


a


M " c ■Sao


& <=>d-<

a


&


c


CM 3 <M


« 


a


S|g


r~


• r« ^ r


W



ci o u


b


5 o~ S o


3^ "

o M

1-J



2« 2


J


g03 O goa




~H -»# O






p pp



~









>> >>>>



> 2


p Jl


JT"


- '- ~

2 EH


p


t




o o o




a c


p


M bD SJJ


3



o


c


a ae


O





o o o




is i£


1


s ss


&


-3 « a c a" <^ © >>

•2 6 B 8 ° a


c a


• s §sl


•g .S A

ai.a

ol.fi Ooo w

V u « TO






S


O <M^


"o o o

ft ft—S c


S. gjL.a


w


o II 5

£xlJ


c

-* >


51-23 £ a o


3 c3 +i u



X


X


M


M





b



fri




t>







E





t





O




b


b>


a






2






N






H






h






O


5



b


b


H





00


31











c






z






o2S

J PS ►,.

gssg


if bf


M to M



£8


m O o?



I--O0




MIN



—I r-H



a






z






<









ci


ci










C3


« 






















S






CS CP


C


E


E


a



o


o


"5



_ci


ci


[C





"a



C3



o


©





a


a



c


c


o


o .



03



j


J3



>,



o


_o



-C






H


h


H


H '


E dE a


h H


2.S


a a


a


a


ci »o


to


CO *° to

CO CD


Cfi


a co

^CM_J . . O


"o







a


o


a 1 ^ .


tSJ





O - in



n a

a— .•

^ffi He


Cu

a~


a

X

H



%





s'S' -1


gco-g a-g a- s a


02 »-,


03


•S


M IS!


SI

















o o









W) be


be







a a



a




o o


o




-c





a a


a







£ &


is


tS


W ffi


w


_3


_e



■bA

•a a



X


X



a c



K


X



a-cl '^ E



bO


bo



£ o —



a


&



ft-g^



a m


a b '

.a o



a«« 








X


X



X






■b






CN O






M CM





















"bo






CNIOl











bo






CM<M











E




b& b£



bIU°



ft2


M O



ao _



ai O



ol Oo CJ




CO^








MINWC)















3




























3


a















a


3



•a

o




fi















3








u








n


cS



>>



J



i-J




ft.



CHROMOSOME NUMBERS IN THE METAZOA


35


tC '"



ft



•*?



to


H





«,-"*! o




ft










rd a



SI


'" H


j"




-3 o3« 





!-» — ;



W


pq


. . o









D O^



ago


S3N



t-s


pq




C5



CO(M



o


o


O-H











3


a




o


O


o o









00



=p


&


3 s


=




n a>





aflfl


S


IN




3

5


■rt a>



to p X =

separa ploid, haploi to po



X



N











o-d^





~2* ®



■b







'b.aS



- 1



N-«C









. |S M









rt





'b '



■fa







'fa



"fa











bO bl) 3



bO M


ago



as




CO O




m-*





3



3







©



03







a





03



cs















O



o



A



J5















>,



>.



p4



Ph



— '


-•


o



-•



~


O



a


d


05


Oi



ft






0.


la


IS



■*



cr>





Ph



d


Ph

^3



ft 3i W5


ft


CO


ft"


ft «5


ft

CO


CO



0J

3


0)


d^J


30


ID

S d


CD


0" r


00


,3

ft


ft


-3 O.


<


=>-<


-•'3


J


^o


<


^"3


— r



• r 1—


S


— r


O




^N-J


3




3


f~<


3S


3


3


s



r „;



m


• r



■r* a>


n



pq


pq


pq



5


1 Hi 5


"c



g03 g-5


"o


o M 1-3


^2 =w


~o


3


^co

^5


H


H


<


S


H


H


<


S


P3


S



as


o













p


p


~



p










>


h


t^



>.


>.


>,








M





u



t~4








a


01


cu


o



c


O




C»« 


Ci


C7i


C-1


£


s


s


^


s


5


g



^H


O-H


p


O



c


o


o



o



5 '








bl


M


u


CD


M


C

tc




--' C3



O


cT



s o




1

s




3


3


3 3


3


3



C



03 Ph


3


3


03

Ph


>>

03


03 03 Pndi


>>

03

Pm


>> 09

Pi


e3

Ph






■5




3 ai



"3 3 m


"d


3 m


3 co









<N -m






!M «J






GO




a C



- O


c


N 3

3 O


rl S 3 <n






" ^




• 1


.

»S


Lo


'LoS


• fl






la

O - "

■5 co




^,2 "3 "3


"3


3 « 


"3


3 ffl


S J3

"3"oj








ft.ra



&OI



a<M


ftrj.








||


O


2 11



oil


O ||

















a .JS

CS




><H


>H


l^« 


><


^« 


><X






CO




X


X


X


X


X


X









'fa



=5



■b


•fa









10


"fa



'fa


H)

1<









^^



^ H



"^


1-1




























■fa





b




■fa


'b


'b


'b


■fa


"fa


CO





CO




CO




cc


1C


>o















"t




■fa


'b




■fa


^


■fa


'b


■fa


■b


ce




s


CO




CO


CO


CO


1"


»o


























S3




M







M !=C


bO


M



M bd


bo


ft







o-g


0.


ft



o-S


ft









CO O




"fao


CO O


CO


O








too


■<)<


CO


-.0 to


t^oo


»o










Cl


CM


CM CN


CMC!



||





II







CO











03





03



3 M 3


ad






03 "ft'g




c

h 1


O





'a


_ft

C9


'3 M 3 03


'3 M

>*


3



03

-a


03 C





3S




3


3

_3



6

a



3


'e


1




n-3




S


'C


3

-a


'3.

CO


CO





1


0! 03 — oj O 3





0)

|


IS



_o


.2


a 2





— c


■"Si





'3.


5


.£•


"


a


<;




<


<





<


<


CJ


Q


fc



0



36


ETHEL BROWNE HARVEY




-j




d


p.


O)



d


d



j3 .a


O





OS




P* PL,


£


CO


CO


" '

rH CO


CO





CM


CM


d


■ <M


CM


H



P.





a



o

la m

« 


£ ft£ d


CO


-a


p,


S



,a'



_i


C-. O


O


"3 P3


T U



H

PS


CM CM


"3


3


S


PQ


S



2 c 2 o 5 o 5 °


"o

3



1-5


"3

3


"3 a£?

a 5




1-H tO









O O








PS


". ~„








H


>> >>








>

w


Fh tt





Oi




a


O



CM


o


OS CM


CM


s s


p


r

r*


'


Pr"


r*


OD


o o





CD




n


SO M



V


CD


a


CU



o


i3 a o o


a


B


a


>>


a a


a



>>


>,


>.


o3


>>>>


>>



CO


03


03


Ph


oj c3


oS



Ch


Pm


Ph


PhPh


Ph




t3


t3


t3


it


T3


X)




a m


C! a,


a


« ai


a to


a co





CM -^



CM -»^


CM +J


CM +i




a a)


a g


_a


a v


a g


a cu


3



£


«B


CO

o


•;s


'c^S


»fl


PS



-3©


S_cu 'o'cu


"3 "3


"3 "3


iu_o

"3 "3


a

H K



P.CO


P. CO


p.


P.CN


Pco


PlO



O ||


° II


o


O ||


II


O ||




-^ II


-M







><X


>HM


>H


>H><


$*x


><X




X


M


X


X


X


X


a



■b


%



"b


■b


■b



m




■^i


10


1^

i



CM


CO



CO


«

^









H


















'b


b


■b


"0


■b


'b





ID




CO


00


















B









<N









H









O










'b


■b


■b


b


'b


■b




IQ


CO


CM


10


CO


00


fH









0)

















c









fc









£ S *

w o









M


bu bfl


bD bj)



to


bJD bD



&


&8


8-8



p.


&8



CO


CO 00


ceo




000



CM


CM CM


CM CO



CM


CM CO



a









g









5











































a









a





a









3


3



cj






C3


e3


a

o


a o


3 a





X rt S «  CU g g CD

"p. 3 <".&


u

W fa




_a "3


CD P.


a

03




09


3

."2


."2


CD

-o o


a


CS

a


S'a'cla


>>



'3


'3



a


c3 c3 c3 c3


03



_o


o



c


cp a cu cu


O






CU



a_a_a a


a



£



fi



Ph



(3


a


raccc


oca


S





^




02


x



Ph






05 CM


,l>


P,CU


■doo' cj>o


a ft


< m


CM




oj:


S


OCL,


a« 


Pw


H



CO


O







>>


>v





CD


£


E





bD





a


a







r^i



^



1-C



a



.*H



CD





O



P



O







■b



1^



CUTS


cu


« a §


a

03


==M


cc



a








'b


?H ^




■b



t^








03



>



0!









M



a





H




?X;o


"3 <" cu-2 Sag




IS UUJSw


&X




CHROMOSOME NUMBERS IN THE METAZOA


37


ji —


-ft


.fl .ft



ft


.3



ft


..ft


3 ft


Xt


ft


3


ft


3




3 »


-Q j3


3 „C


3


3 3




3




3


3 es"


3



3



3




3 _J


3 3


Ph « 


Ph


Ph Ph



.


Ph



"1


Ph


Ph "


Ph


co


Ph


CO


Ph




Ph ^


Ph Ph



























"co °


-^


to to



o o


to




-^


« 3


'co


O O


"co


O

G


« 




to O


-»-> ^>


a N






N




N


a


a n


a


N


3


N!


d




5 N


d d







KH













t-H


>"H 1— 1


— ft

CD X


cd"


. a> ' - ' .2


-H


ft


£



cd"


_ ft

CD" X


CD"


ft


CD"


ft





CD M


Qjt-H gjl^


cd "


M ft

'3) ft '3l CD " O


£


H


"3>


i


w


'cfft


CD "


'Sfo

H


'3 n

CD M


w


'S


ft



'§ft H


bC ft it ft


^-J^IS


c _r


a - a



t."" 5




l*"" 5 c



1 - iJ l -'»


d

C™


a

u;"o E




a - ^"5


d - d

O 1-5


ss


e3 "5 03


a


32 O


03


s


3;


- cj"5



ss


3^3



- "

03 "O


9 :

1-3


-1 s


10




03-O 03" 5


o


o o



o



O



O


O




O O


P O


CO


CO CO



OS


CO




CO


CO O


co


02


CO


c^


CO




CO Ol



S CO


O


O O



o


o






O O





O








O O


ir cd

^ s









CO


CO



3 CO



CO


TO



CO




CO CO




e


3



d


a



a


a


a



d


a


p


a




d d


d d


o


CD CD



CD


CD



CD


CD


CD CD


CD



CD


CD





CD CD


CD CD


>


> >




>



>


>


> >


>


>


>


>


>




> >


> >


ID


© CD




O




CD


CD CD



CD


CD


CD





CD CD


CD CD


<» co





+i














J -«J


co


CO CO



CO


CO



X


CO


CO CO


CO


CO


CO


CO


CO




CO CO


CO CO


sj




tn


2



i


d







.j a

co^O



O



■a >>

C3-=



M




.2 2




_B




_c






2




a

03



3


a

•—


ft


1

M




„. o











a


...




tr



~


ft






CD CD





CD










CD CD



b a



>




o3




"3




o




o







"3







a


13




ft

sg




ft


s


p


ft


=


OS

a




"3


ft

S


O


CD 3

•a -a



■9 d


s


CD

u

CD





o o



o


r



o







2 °




n





9



CD




~£.S





in ■H


C


43



d




3


ii


3


c3 ft



d


31


03 St




X




X




X






!*



X



S



O




|S














a a

























c


























(h
























ft ft

























bo

























10 10
















b




b




b






b

CO


b






.




•^



IC




o



>o






u



00









CO












































O










b


b


b b




b





b


b 03



03 ft



r ^: J




b


b


00


Cd


cc


CO









CC



-* ft




(



■>»




OO















00



00

























t^




















u


























a













































O







c





M^



"t














53







|





ft 03



r














ft










io ft



ft













CM











0000



oc




















~Z c


M

CD





























_;
















.C




ii :








CD






"od











03











iS






3

ft








J>>


h



o




os :

■" 03

o'S

- CO

T3"3

"3 o

it bfl








3












"3 o

o

0) 0)


ft 03 ft

CD

13,


a

>

\


-a o

u

a

CD

-a






1

c

b


1


1

.

■1


CD

1



■a

CD

1




CD CD




E

a

c



c ( Jt

$

C


-0

CD

03 O








. 03




















-d


2



IS




ISIS








s



ISI







T



~


"S


























3


13


'2


IS




15 "3




in



IS



la



J3 C




~




IE



'2


Ic


ft


c.



ft




o. m




c


1


c


i


0.



ft a




ft




z


.


c


a


<J


<C


<


<




<




<



<■



<



<




<!




<



<


<


38


ETHEL BROWNE HARVEY



43


„ r


ft i


a


43


43


o


<A J5

SP-i ft


ft


43


C5


rff ft


J



3 P4


3 - 3

P-t —


«> K


3 Ph


3

CM


a


C1 ft,


^


a

Ph c=>


a









■ "O -


a




a



— ' a





o C<o






CD

Z


"io


.J ° -J co O co


o


CO


■g


CM


O 03 .


O co


CO


-J o

co O


■g


a


a


3 N


a


a



fti ■>-> C


N C



a N



PS



h^ _ t— 1



t— i




5 CD ^ S ci

ScO!







© ^*


a a =a


CD"*


w


s

a O


a .

X CD rt


~z


— ft

CD" 1 X


.S*^


a K


M q. CD u '


'So gK '51


a^ n


'§>&



W '|ft


N


'gft W


'3j ^

0) •—



a cd" 3


o £ o


5 3S O


a §3



» is


gsaa


'o


O >-5


as



o


O


o


pq


S


<J


u








S




OS




K

a >

as



r? IO CO

C5 O


CO


-

>

o



>C CO


_=


w C5


CO


p


o o o


O —1


O



cd


o


o o


ft


p o


p


a to «  o


a


CO CO to

. a a a


fl 3


C


B



-t->


a a


cd

pq


a a


8


CD


GOO


CD CD


CD


CD


^3



CD CD


o o


O


>


> > >•


l> >


!>


>


? 1


> >


q


> >




CD


0) CD CD


cd a>


CD



CO


CD CD


O


CD CD


O





+* -*->





w






CO


03 CO CO


CO CO


CO


CO


CO


CO CO


>


CO CO


M




JS O M.2 fan CD -*J






CO" CD


t- .; bfl a



^ a

n_0



M



.— rt ° H

a^ ^





o


CD 3

OJ

a co


"Pill



CD

a co





o|o8





J2


•5 E


.5 2.3^



CD 2



S

a







a

CD

a

o


43 2"g

ft.fi s

O


-0 o

co cji a

-^ C3 • ■°. Mft^

a ° fi E

CM



"o a

Eg o o


Q



b


"b









5 i





■*









a













H

















"b




o

o


'bo



b



Q



■*









to



Z













cm













a



hD*-*



o


o




o



o





""cd



a


c.




od



cd



'b


'bo'b a



■b

p


o


o


"bo ft


Ot


i^a


o


1


t~


"OIOCIO



-<c*oo


00


-*


o


10»00


=:



Cl














H













»

























O

z




— 1 >>





3





o


3


2 a h o h a

fl, 5 z n ■*■ a

o



CD I*

do a a




"3

a ^



"CD

a * « a


1


a ° "3


1



oo *




0G


Cfl


o


oo


cd





Ci





4


C3

a



an, a

oo •*


a bfl

CD













z







OS







<















2









^








43









a








a <!


.








o





>>

u

o




II









■;£




"a


J3



CD


i— i




cd









bfl




Cd




M









CD



a


J3


u




CD






||




U




>


CO •



43









bfl



o

a o.

00


■a


o 2

a S



o

c

CD



a


a

o


1



CD

cd O




CD

s


CD



o


£j=



o


£


>• C


d



c


b





OCS





t

M








IE


43 CD



J3


4:



A


CO CD

— o


'i


T3

IE


3



a


a=>



a


c


i. C


v a


a ^*



i, a




<


<




<;


<


<


<



<;



<J


<


CHROMOSOME NUMBERS IN THE METAZOA


39


o


=c


co 10


[



^5

3



a

CO


Ph



d

CO




— ! 3

Ph



d

CO


a


d




OS

o

d



^ en

33 CM




p5 "O r," oj


ft

cjT


C.P. CO

d


"3 o S3



a


O

O

SI


ti



"3


s:




a



o

S]


"o o


O S3



-*



CO ^ CO .




■ dco r a oj-s _:

8rA 1


"o o

N



















^3









01


d




d




a






d



a








cj cbcni a d


d


£3^3 ^ . CB

O Cj~,

O t- C3


X



O-

K


CB-H


•s.




o'



X



X



J2



OJS




gag hS ^S


x


o



'§>d

a U


o

"5


'E*d


3~




'Ifd

a O


O

!-5


H

C CO >-5




i-a

o N



O o O t. SJ<1




Scso

o N


OOOC-J

















t^



00 OS






po


£■<



















Op








,_.



to


CB


CO



Cs>




co



OS


~.


C33



p


> .




CO CS 3S C-l


»o


t-t (-.





o



3



3




3



o


O


o



o






o o c; -h



,aj=]




















^3-a






9 CB


















3



CB o






CS cj


cj


c




a >


a >








c

CB

>


a




CB



Cj cj




a a a c


a


fflOK

ace o o o >>>


cj b[



>



5 >




>



3 M


3 SO



3

a



KPQ a a




0j C3 3 3

bd it to bO


3


o



55


cb

55


cb

55



g 02




O

55



02


O


3



cj



o o

»




3 o c o


o






5|




a





T-3


3

r











.SX'bS

o o a • 3

>q-« f, a -3 a-S:







c




t






t^












.5


"3 a



a cb "o p.



.as

,2 8

Qi

a


OS


a


i-cj

S


OS O




s

a


CJ

s


en p



CD

"3



3

a

.5

d











o o







o


D










J • o ~



o



c



-tJ>

a


o


=1


_e





a



o



2






2 « o §."•

O ^-i +j c* —



X



x



X



X






><!





X



O






Q












































3 3






CB bo




















P P






— — -"_ 3



"b



•h
















aa







CO



c~
















'bO





II II *0~




















coco





























COCO

OT3





















■b k



J3



-3


CB













J3


o







^o^


H<m


03


H<n


oj

C-l


CO



UO






5





Hco


d


'bo

CO CO






^^-S £



.J3


0)


•J3


0)














0)






II II '5 a

»— — u CB

COCO r^ fcO



co £


c

cb

Ml


"b.t:

CO £



bO




■*






-»*





'bS


3








of



c<T














CO








02



<D























a



2





o















Dl







c





a















c3





'bOi-Of



■bS



"b



"b fi



--.




o


^<



CO CO CO


o







ira






















JLJLJL

CO CO CO



o

























CO















































WJ tc



bO o


£-5 o


bi



"1
















o




&S



am


•o.


a
















Ti








CO



co a



















coco



m'b


O gj


lO



LOCO















Of





II II



1C CO

















it bf



cj























a o


c3


a







CO

















mo



o




IT5 CO




















CO CO CO


CM















« 


eg











CB









,





' ,



s

e

"3





c

c




CO

Q. (h

cj

n

o c "oj 5

— [




XI

a

3 o o u

to




ao t, -_

Kg

o a

!* cb

2 s


cd

3 3




"3 a

1 '?

3 • M

o ^a a o

'S


°


3

-2

3




'o

15

3

J3

a

03

d

a c3 ^o


"3

3 -3

3 03 X C 3


2 cj

3 u

."§1

« .3

>> a

am

3 3

bC SO

aa

a s


•at?

is

"Sa


3

3

a

CB

cj

>>

u

&

CJ

3

X

_o





a



a 1 *



O,




a


c

a




X




CB « 



<U CB


-a



-G



<:




«C




-sj




<;




<



<




o


,J




s


"


PM^


Oh



^



40


ETHEL BROWNE HARVEY



ft


ft


ft


o


ft ft ft


ft


ft


o 05



|>T


§■>•


t^


CM |^ CM OS


ot-"


t-T


■on



o



o


ftco _ r r o • § o o

3* Q,fO O O


ft-3

si


o

N


ftft

coco


K









w


ft


— ; o.


ft


=cm ft ft ft

— &


ft



fc.


X


3*


X


3.x x x


I&


X


C J^


s is


K


w


B gW W H


W


<N



1-5


• (JOS

O 3 S3


ij OS

£ CO


• C ■ OS t" OS (* »0

•^ O wCO Sf^ «M

2-r 2=3 3* gcM '^ S^ o o o


.2 2?5


o


o >-.









OO OS


K








Op



OS p


«OC5


OS


CD OS OS W lO


to OS


OS


-

Op


O


OpO ;-> rt


OO


p


X. J3


pq








CD CO


a;


(2


a d


c


c 3 a b a


a a


c


« 03


IS



cJ c3


03


03 c3 03 03 03


c3 c3


ci


CQCQ


o



b/i bfl


bD


M MM M M


bl) M


M





C


^- ~ ~ J- S

u. ^


h


3 3



o


O O


O


o o o o o


O O


O


C O



s


ss


§


sss s s


SS


a


»






«^ ft ft





in





_'2.S oi 3





M





-3.0)





PS

a









« 





O js-,** r © C
















<M









M M









M M









o o





Q





cj 03








ftft





H





(DtDOM





fc

H









o 1









Q





tO „ -g CD

.O & fi








ft"





• w





CO












1M


H








a


fri





o





><





c3





o





bo ° CD (O <M




"~" 0J









o g









cq


6


a



a

CO


a


a

IS


a


a


o H o

" K 5

£5 * r < m

a *- o

o


br bD



M b/lg

ft 2

to Oo

ooo


"t^ WE



o



br M 0) o

'bo

CM <M CNCN


u- CD

Mo

m S

CD 3


» . M bD -_ fe 0) M

wobo

M c to to


0 _^ cJl S M S

» bi


o

03 ft



00



co



to


to





mr ^


















cu


3






03




03


XJ










_o





a

3



03



£


H)


a




ft


M



cu








br-

a


CS

>.

H

03


3

C3


ft

co


X

J2


o

JO


3


■g a

i3


H


o



-a



"m


CO


03 03


m


c3


03


a


CJ


03


03


i- i3 3



a


a


o


i)


CJ


o


Oj CU



X




X


X


X


C 3



o


o


_o


o


o


o


o o











">>


">>


">>


■>>


">>


">.


Icls



.3


X


.3


J3


-q


>£)


c o



Cm


a,


PL,


0,



Ph


ft,


coco



ft




-O





■o


3


CO


r«"


-■?


Ph


ft



ft


. CO






OS CM


N


■#


o =■














3


w


33


CU


"o


4"



£2

03

ffi


aq


o


M









fq





-0






t*

n


o














bfj

M


tsn


3


C


a


CU






o —


O


o


CU


pq


pq


X


cc



•S *>





« ft





.Sc^



5


3




3


CD




o


o



o


ft


-2 °



ft


O




■*"*


p-,


o



X


X



« 



o


b







,-H



^ H



O



rt *






b


b'b



b






^


^- *-^



^^



o —



rt






-b


b'fa



'b



^h c;











8



bC



M M co



a



» ol b





OO'J'OO



tM



CM<M<M



A



3







M




c o




3













t~> ^-v



-O eu





03


03hJ





03


3




£*


ft


C"




^ ft



03 ||










O


X —



3

ft O


2-S^

•= 03 5,



C co


ft




ft—


<c


<;




<:



CHROMOSOME NUMBERS IN THE METAZOA


41


a




a







CO



Oi


CO



2





22


CO


■*


CO



-*





CO













"



a



- r d





d


d


o


a



"b







o



-r


.S =»





o


•*


N


1-1



N CM





CM


CM


X


[3


"5


a -J

r*5 3





"s


3


W


"C


rt


H « 





P3


pq


i^ca


O


^ H

tiS — <■





_j


^


•-5


03



2^ .2





o


o


»-)


2





3


5






it





to







o





o







Pn





fa




>o




-a





73





CO


r—






CO


p


r"






^2



M


>>

o


cj


cJ m"





m"


M


a


S


.? a -'





.S c


_c











t-,


u


^


'£* u o







o


c3


o


o o —





o —


o


pa


o


PQ


pq a




m


pq















3 go
















a



13


.? c





a


a














©


a> a>





o


4>












o




"o o





o


o


a



a


a a





a


a


o



o


o o





o


o









-♦^



X



« 


X X





X


X


b




b







r ~"




CO







CO




r^r

















b




b





b

W3



rH




00





'~ H


, ~ l


CO




t>






■*


T-T












b


s


b


b b





b


b


■*




OO l^





IO


<N



to



















S2





M





M


M tJl°, 8





a





a


O- 2




y







<n Ofjo





if






C5


cotco-*











CM CM CM CM


C3

















03






"S






3


=-.






o


2


.2




M O

a-a






a

73 C3 3 CT c3


09

2 a

03


3

2

3

a

0Q


oi

C


go

3

o u

ft g


o3 a

c3.c 3 a


3

a

bj


D


c



3 'u

03

a

3

a


O








-.

d


GO


co



.a


-C






sj


3


3



a


a


P


a


p



T3


a


a


o


c


o


c


o


o



CD


o


t-l









03


03


a ■


"a


J3

a



03


a



3 fj


12


la


<


<


<



3


J




PU



a,



Wilcox, '95 Bull. Mus.Comp.Zool. Harvard, 27, p. 3




33 03

73

-3

03

a '6



CO




"53 b

to



60

a



B a

e

Jjj

03

a] e

a




io iC





o o







te to



o o





o o



o o



N N



o o





>CM >t




-C .-C .



gaga



<l <



CM >1<











a c







a a



c3 03





<U <D





Pm Ph


a



-i





u



CJ





U



■*




bo



CM CM



bo



CM CM



if U:



ago














-c







3



a



(8



>,



u



o



a


a



ft








■*


•«»<



■*














o






o



o



N


N



N



ft


a



ft



X


X



X



H


H



W



si


t!C»



t*CJ>








1-5


i-a



i-a








O


p



p








60


M



tr



a


a



c














o





PQ


pq



CQ








CO


03










a


a



a



"<D


o










O


o



o



a


ft



ft



o


o










X


« 



M


ca

a


b


b



b


CO


CO



■^




















bl











3






US











b


b



b



CO






1-4


1-1



^ h



IM


(M










b


b



b



CO














a



M




mS



a




o.

COO



m






CM




CS CM






-; — •



03


c3








c

CD



03


03




s


^j


.5



>


>










J2


^3


p





c3


03


03



a

OJ






CO


c.


CD








a



ft CO





















0) 2 O °





c



<


P

PhP.


42


ETHEL BROWNE HARVEY



a


a


a


a








■*


■»<


■**■


■*







£3






O


o


o


o


o


55


o


o


o


o


W


ISJ


S3


S3


S3


PS






w


d


a


a



fc


X


X


X


X


IS


H


W


H


S



3. ^



P


.;o> 2*



O


o


o


o



>"5


i-s


1-3


1-5


« 






a










r~


K


O


o


O


o


H




"



CQ


b5


bi


bE


ex


o


C




a








u




t*



o


o


o


o



« 


e


PQ


P3




"S c






JS o







-»J








M


^ H


•3 a>


rH


H

PS




a


a


•<!



-2 o




s



o a




a


"o


- M —


~o


"o


M


a o



o


a o




h.2


43


■^



X


M


X!


X


a


s


5b


■b


b


H



f-ioa



00







o


ceo



t-~







H






H






>

b


5b


b

CM


b



^ H


— * C5


-"■


oo


a


o


ooo


_r


t>."


£






CM






B






H






o


b>


'b'b


b


■b




■-HO



O0







H






CO











O






7,






o a o

h S h O H a







bt


SX


ho



a


a


a


J « S

£^a

Q P.







~


CO CM


• o







2






<:








-^"7


s




"5


is


_o a





cs


"3-T3


S"o


a


co

a 3


"3 a

3

Cp

3 5


8e

al —


o


a

p.


bt C

03


.S.H


"ol 03


c


co


j £»

a a


CO




0) 0)



3








cS


o o


o p




3


u u


u L

a




_o o


•o-o


o



03



CD CD




bo


SIS



2



«!


oo


5i5


ft.



a a


a


a


5 d^"


5 a-*








.


CM g


CM


N £


CM



->•


•*


^



■*


— O





^*






r


a o


T3


a6



.












o


c


o


CM ^


03



>


o


o


c



o SI


1 3


>


t 3


o SJ







uJS


03



3



a


a


ft





33



X


X


X



X







W


H


K



H


N 3

• PQ

■3


— :rj"


S3 3


— i-S"


w


i


OS ,^(35

o

1-5


=


— s


»!C35 1-5


OCM

o .

si a


_jPQ OCM

1 on a


o^ i-s







«*











r^



r^









c



c




r~





t^







~





p






o







3



3









cS



C3




hi)


ti


bl






H


c


.s




.a


"3



"5



a


c

s









o


o




c


o



o



o


PQ


pa ■


pq



P|


M



Zx



a







-a


•*j






+:









,2




O)


_B



.3>!





c




a


CD



a>"S



£


a


CD


£



CD


"o



og



CD


"o


"o


"o



■o


a



&- S


CD


"o


a


a


a


o



03


T3



o


o


c



o




-28


>


o





-*^


^H



o


— 3



X


X


X



X


M



X



X


b> b


b


■b




b>



b



o




o


b>



o





^ H




- H


o



~"



1-1


c


oT


c



oT




Oi



o"



















ojoT


.










O "-<


?



b


b>





CD 3^.


■3


5



o




b


t










o


o



■bl











° c




C35


ca






2.2

oT


o 2

c a


o


b b


b


b


b)



b



b



O




o


o



o


















^











CD











CD








M




M« 


bt} bt



be ex



M



a




a"

<° CM


a 8



&8



ft



C33




S.o


oo



c»o





1 - <




CM CM



-H CM



CM






o











S













































II *^






oj







^ 03






>




03

c3




03 tj -^> 3






kt







03 CD





e


3











(

o


1

s


3 Z


2


O

3


o 3




5


1

c



3.^


a "C a

tr B

b CD


13

cd

a


3

33

a



03 CD

H 9" J s




c c

>


>>

|


J3-C c« oi

CO IT

cd a

L- 1 CD


o a eg

,c o a


o

a

CD -3 CD





.2


5


o


oc


U


W


H



w



a



T3 Jj 2


CHROMOSOME NUMBERS IN THE METAZOA


43


-* ft"?!


p,


<M P


^





■*


Q

-T






C"> £


o3


o


rt 3


SI


"3s


w


ft


SI 3 .« 



H




g o

<


O .

S) ft


dS


■*




r 1




c




3










d












m





.-I



-H

a



d





oi







o




ft



ft


o



o





M



X




b




OS




00




b




Os




OS


b



b




OS





Ml hf



M


°-2



ft


m O



71





CN (M




a:












oi 03

E



d

03 03


.O




03



3




-c






CO

10


oo



OS


o"






ft


ooo

OS CO


3


3"

en


os

CD



3




"


ft











u N

•*-H




Ti^lM




lf3 ^




ft


ft



ft"S





<!■*


ftft


P-1


ft


O ftft






«*




.a







S:3










1







s'l


OS-*

«eo


SS


rt


t; 1 ^



O 03




X!


X! u



2-« 





S"d

lis


n: n


"3


•^


I«S23




<^T^




x

03 Id

"c o


X, 1-3

"o

o



a- d

i4

d Ih


<* ft




d d « 

c3 03

d d


OSftN

• Jd CM J5


i^^2Sl

• 2-d

L-rr. ai "^ h

P5 <




03 g

2 ° n




SI


SI



<J<U





a,


« 


IB


<i




ms




_







&











-d c;


-d

u







ft o O


COt*


<M


CO


" P














op










's


%



CO lr~





-Ojo


-

J


s


01 01




o« 




to


<J)



OO





dP


01 01










M


J2







03




jd


3 d




03 03




a d


d p z.


<5


> > oi o





bJ2


03 03 O 01

d d o o


2

03


o3


a 2




"c8 03

d d 03 03




Ph


&







S


QQ


»


Z


< <




OO



-d-d >>

111

Co

« s

a .&

W) "J


3


ft'*


sd


ge

S2

-S-d I

t-X






3 s -d


a


"ft

<

_d


>


St

03




O o d o


X 71 ftS

"o

J3


d


In 03

"3 S


p


ft

a


>>

3


>

o

St 03


01

> o

St

03


O

.o

03



ft o3 d

TlJd Cj

a ft ■


d-d

jd ft'C





csi o



O c-l




^


71

<3



ja o3"0


QQ


s





Q




W





<



O




<





























01 -^


















— ' d


















S o









o




b






b


O g T3.S


d


b




b


o

e


03 03

■a


oo




CO






CO


a


00




CO


'a











boo






1


<











CD 00 CO






s


































us



a




b





b


■b


03 T3 "O 03 >>>>

ss






b




00




CD





CO







CO













b^

CD 00

















^

















'a?



















7i -a

















3


"O 03









o




b





b


d o


03 fc.



b




b




cc




CD





CD


■a



CO




CD












b

oo


bo

CD 00


































^~ 01






.5.2

3



,









-23



S






03

ft



M ft 71






or

o o



M 55




mS




CO




CD






CD


b« M


1^


ft b




cd a>."













CO CD


o3

ft«w


71 O CD C-l y* CO




^11













" ' H


r^CO














d















































W



































« 












II






























oi




s




o3 : „




03





03



03


03





"o




cs a




o





o




01



d







cd c 2 a




s




td„


cd


cd



u




'>

















03




^S



















CD





01 01


01


o






ft




S:




s





a ai


E


s



_03




O 01




.2 II




m





.2 >s


2


2



s




_o




ft—




'ft





0,01


'ft


'ft





>i




<!









<!



<



<!




c





X



,5 cm 3 ft

tCco


a 6


d d

aw


44


ETHEL BROWNE HARVEY



o



~H °"



"cm T 1 *

CM i", CO



S



.CO


d*A


"o


d


««T«S*


Z H


o



co r


"o--"o.-r o 2 o o


K




o.

tS3V;N« 


H

K


co

'S2


<!





■£&


H->

C3

G


g 2-* ^3 o

<!l-5


__; bC_a bO



tS3


<!


pa pq


IS






H



"*



oo >c


>


-*



^iiO


O -H


K





w










en




t- u


a


a


CO

0)


a> o



o



a c


o> cd



u


>


fat! bt


a- £



o3



CD CD




s


8


kw


CO CO





_^






03






s



CO






M




o



« 




-Q



-<!






S




%S





a






>>'S,


,Q





c3 to


d





s



a


o





H 1


-*





a






H






rH



Of




O

1


o


7 o




a






fc









<s



H




a



Eh





o





o



O


o


o


£ o


T*


-*



O





CO



Eh




o


03 S


6






&






Q a S






° fe §






a IS ►,






o P.






Q






Z






<















CD






a


c3






o






"3



la


0)



co


o


>.



CD



H



3



^ _



5


3


,Q


bl


,£3.2


w



03


03



PL





"o


in


'ft


5 o


3


3^



rt



o


"oS a



"o



2


D. o



be


3


'a o


OIL



<


o


O


Pu,


















pq



pq







o.



CO


00



c








d


J



cj








d



00 CM



X



o

X 1







Zool 35,




>,



>>











o









CO



pq


P4


X


pel


CO

t





.2 A





X



^<









"o


o



z.


s


a.






3 N















ua


p



~







CM




c



c








p




a



s










c

03


03



4J

5







.2 d



s


u



^









c o









1 2



u


Q



2







W CQ



1

>>


CM


a

a


>>


M



7







o

II


^O


o



jt


09


»,


.a G


a

o





CD

a



-a


s -a


"o


"3 « 


u

a3 o





bl

c


c


a.


o


ja



a


o

a





-, o


a




EC








c w Q. *

to a o


CD


a >


'S


Of

CS

a:


d

bO

a


a

CD


o

a

CO


-G





-*


PM






X

















Of


IS S3



o






■b


c*





"2



o






oo




i












o


'1

























c













>>













u













6












o



■b


o






^o




O ^



CI


3






o




O













o













J3













O













Lh













&




o







o



O "-'




c







o



01 O

























"-ICM













1 -H













o













^












oo tn „
















03


a

a


93


5 c



a

— . o

bitr° m

a oo o


J_


O -O

O-JTS



°2!


a



a




CO


o


'l 1


cjo -h 03^"




ooc




oocoo


acN a n



CM CO





-ri




CM




11












a


CM


















cc


,





















,J





93










o

CJ

c3





3


tH









G



















CO














C3





s


CD








M






c3



cS






a





c


O



a










F

t-i






a


cd







J3





i


a r






O



0.












O





o






T


"O



>>





a


II .




C


O



u





s


^> c






J3



Q





fc






pc


ft









a



CO




0.







CD




CM







-O


N






J3







>-!


&



"o

O


H-i



N


tSJ



00


CM










bO




G









A


G



CO


M















TJ




G




03







Ph.



93





•*









a













ta











Of




■*







Of


Of



CM







O








03 U



0^2


°-« 






CM-*




03 03


COO



O O


r— CM








CD

















3





bO






M



cj


bl




G









a


3







a



&



►j



CHROMOSOME NUMBERS IN THE METAZOA


45



y _ t



w _ i



-Hrf


©




J" — '



^r





COO


c






« 



•o


OH


GQ




CO "



d


d -j


ftft


~


r-

CS1

33


d o





£t






JS



d <*



d


PL)


o> —


HH


d


d


Ph 5






-O


t* • •


d



J2c-5 r



a



DC

Is


6

s

C


dSS


0<MOO

^d£


ro

J)

©


£2 M

S3 ■

a d« 



1-5



1-j


c o


C>T'-5 > -5


9Z rt


s

o


%&*



o



d


fi


"doS,


fc


o X


£ ci



CO



CO


■*


COI--O0


oo


CO co



a



o


a


OOO


o



o o



^




t*


^ Ch



■~


"


- u



a





0)


o



oj


0) OJ







-M -u



+3









cs


§ it


71


S


A


« c§



o



e


©


o o


©


©


©


© ©



a



C


a


g g


g


a


G


a a





o


o


o o


o


o


o


o o



P



P


a


ppp


pp


P P






l


i s










o


a .2










S

ft -3







o



o


K


cj





©






to © ©







a



o







09



c



c


_>





G



V



01



ID





OJ



bid


t>n




M



O



o






o



c



c


O


O r!





G



a



a


Q/ C a


o




ID







■-SfipH


g













CO



Ph



Ph


"






Ph


01











S











-a





















a





















-o











oj











u









































a











(B











H



•*









oi


Is











a












oc










fh


d









»-g!


c









CC


— i










t~



















_>,





















3






^o










00 00





6

0.






cj


OS










©





CC






M










ft to





"o






CC






03











0.






a






00






c
















a











3





















o





cr


s





S





3


o






_3













J



c



.4






c3



(S




t.






s



>



2


|






O



f








_o













c



£ a

Sz







'S 9

o

Ph



<!

<


p. 571 Proc. Amer. Acad. Arts


CO

ft

CO

-a

c

C3



Meves and Duesberg, '08

Mark and Copeland, '07



1st spc rudimentary with no chroms


o

03

~

'S

ID >






+ s

CO CO





c

U

e

ct


>


e

s

"5

c

B

E

ct

, [

>


,





Q



ft




d







r









•CO

ft .


"oo



"co




a "






>« §


2 m



- 8 !2 on










N 3


- 3



- 3




a-g









S 53




—'JS


rtj


( ;













c





tS3<J


N<!


l

tsi-<




Tj<lO


M«lO



Tt*»« 












>ra







CC














II




& ft


ft IS


>







OJ ©




c


c c



C G










<U ID


ID OJ


r.

o





MM


^W


m!^


a







fe







=.


x a












a.


O) J-j






'-^











90



^








■b


'b




"b



oo


b










CO


o

CO


-r


CO


















OJ

















^2








ct


§




CC


'


OJ

-D OJ

J3


ccf


5












c

a







a










<



<



>_


tr



46


ETHEL BROWNE HARVEY





a




coio


r~




Si

co_J"



a o


ft*




Z








a


■a- .


•* l


K



- 3



K b Ed K


IS


57=




_• a


— —




o o


o„




c

<



t>3-<


scar






>





M

a


M -*


iS N






go


- *


VS



n

o



a a c








'53 '3 MOD


,-.« «§




« 

s


in 0" + 3Y i oid. X poles in lits in t anaph:



-c



XX-E2&5



CO



X



3





a













a










9


Q


C5



I


B


"b .




OOOO



Q



CM CM



O





P4





o a








a


H Eh


O




O 1


b^ '


b




OOOO


CQ



a


C<I<M



~


z

cm





H





H









o


"bo


b




OOOO





<M Ci


CO



u





on













Z


fcfl "v




D H g S W H

si K z


& g

CO ° °*

""bog




£: ^ 3

O

Xooo




z


C. fe



<!






ci





Ed





O


a




•a


a



0D


3


•3

a



o


C8



a

a.


O


E



a



cj




OS


£




60





3





B





£


a




c


co



-^


>>?

CM.* > O



a ■ cj a

Hi .


OCM

~ a



"5


3

t-T .00


-"


CM


cm"



ts


nz., 1 Univ, Agr. . 3, p


Oft


N oc

~z a c » NX



1


< .



c


— i


-J 2"



,

1

Zool.

Imp.

Col


^j


=. d


O CO

p 3





a « 


a <»



s



<


<



c


C5 Ol









CO


CO





r 1


r 1




HE

ci cS



a"


a"



a


>>>>






4


O O



oS


C3



a


HH



>


>








^_,













CM







a -5







O CD













— 2







C.







-C













cS







a







■b



b




■*



00



'8





CM



>.







£







a







3













ci









b



b


b




00



OO


h



CM



CM


IM









1


OO







CM







1 CO



b


b



"c




OO CM


t^




b






c


00







CM







DO



to 00





0.



Q-g





m



CO O





00



00





CM



COCO





co



n





CM



10 10








.2



C3

C








'C



.*








cj










>



■3








t^.



a










C3








'



E"








" X


c


O



O



>.



£


£



s



«£




X



X



•a c



c


I >>



>.



-Spa




J3



-^'m


a 1 *"



E


£



£.2


g II



c




p*>


J3 —



a

c:




oa



r- 1




. Soc.

p. 179 p. 189


-'


^


^


a




a


a


CO ^

rt oo









5 -* n "

Dh n x


to"


«*


•<><


a *

to






CJ


>2 an



Camb. p. 44

Geneti Geneti

Genet nd 229


S

a


a

O


.3

< 2



o„c^ ^


U ~


«h


b


— J3



CIS 3 2 3„


3


a


O ° —



t> OOO ^h ^ l -;'-o


O

i-s


•-a


O >


O ~* CM CO


«*


■*


■*«5





















tn U l-t


Lh


u





O O 0)


a)



a


^> *j



X CO CO


0]


a


cj


CO CO



6 00



c



a a



a a a



3


a



00


O


O


c




Q PQQ



2


MM'



3311







'S'OCM













a a












-OT3"







V







bo's







S C c







•— »«H #J







XXXo







X












i



oc






b


°*.





'fi


00 00







CM CM












5



















O


'" a


10





CO


4- co


_a






<".£ a


co'

5 c *


3


b




o.« **


5 y


"S





b'llo


eo

CO





00 00


r





CM CM


CJ






^_^







1^







Cl







©




Of




C3





b





co 1 10 O







CO






>o




bo







OOOO







CMC!







--^ cu







CO c

"5 a




bO




(50ns




a-7

CO +






CMO




M Mm'S




— O




a £







m ° o-£




c3 O




co co »*3







UO W5 lO











C3



-O




>.




5 c3




"3



cj



Is




J^



3



h




.3



a










S IS




'p.



K






03



X



CO


w;




ta

39



>>

4;



11


n 5 S




a

03



J2A


u



>>'co


3



« 




z



O


CHROMOSOME NUMBERS IN THE METAZOA


47



t^


a



a


a



co














30



00


00



O,


j


^ai"


r



"5


5

HO


v.


b.

fro


CI




3




3









|


03

z


3


z


03 Z



„•


a


_


ja


-a



8



o fr-~*


o — 5



■«1<iO


•* >o


»o



























&


E


S


■s


'$






CD




B


c


3


B


a.












0)




w


M



W


s







T)













a







S







ft













$












i-l







•>*










b






C







CC





b



b


b





e





co



ec



co



03





a>









c





a









cfl



C



fro



o

a



C



c3



o

.a o



F



03







O















£





83







c3



O



_



S



Q


O.


IC









-f




^*




a



a





CO









c






X.







"3


"a




£


CSS


N


N




X



«-^




~" fr

fro


fro



LS


<j


<J


<



o;







o^


o^


-a<








3












fro


fr



C


o


1













W


co


co


CO




M
















&






o






>>






fro






JO
















» to











a o






O









03



b




S
















a]






a











j






m



co





o





+ b»


o o



e<



1


o




CO CO



CO









c






CD






C5











03 fl



a






E


OU

'42 ' o






c




fro fro


p


o



13 fro










G


as


>


» >.





fr

fro



M


fr-lfr-o


_


O


o'b


3 g 2 o 3_i

r +3



.g



a



a



CO .Jos


CO ^ ^00


T ~*oo



^ a


« §)


2 M







-s« 


-3


- 3







^C3


S o3


8 03







U-S


_ IfrC


r='-=



a*


° P-.




C-0


N<


N3<;




■*-»


oOOUT;



























fr


=£ &


s &





a. cd



_-o


fi B


B E









CD CD



o


fr-ofr-i


WW


(I)

















3









o





O





Z





i--.







■b


b




OS


CO



b


b


b



o






<M


CO





a


















3


fro






3



a

03


03


a

3



03



"*











>.



fi •



a





O


2




o





<



<


frj




C o

O fr


■%%


48


ETHEL BROWNE HARVEY




a


3


"SS



3




00




T




s



.fl-H



— > ,fl-*



Z,

m a


o

hi H

a


.OS


«co"

i"3


<


ft <ft



a



S|


~


- —


t.


03


•S-S


c


u *~* u



PS.



IB


CO, fci







•S>


'2


CD ii»

> •->

cd







o





a


N



CO





a



>>



>>



OQ



_o


u


CD


CD



2




o





o


C


o


c


9


CD




u


■a



~






o



a


O




M


ft


ft


ft


ft






>>




>>





a




a





c3




03



w








M



_c




C



K








«ej








S








K






a



PS



2 ©

£ 2




£ CD

CD 3





— . Fh




•-I ~




P




Q


3 C





















hj






















|


ft







■k


H







Q








o








ft








O














1=


H Eh







ft




■b




ft


O 1



■b


•*


■b


<




o


1


CT


CO





CO


CM


©


CM



a







M


2








H



cd






H








Jh



°rt






O



— -co


■b


b






o


OS


CO





0)


CO


D


CM



BO



O -p CO






o








2

Q a o


C.





M



09





ft



3 & « 

Sja

ft B, o

a Z


o

CO

§





+ o









■<









d




a







CD




"3






H


jg



_T





o a

Pk to


cd

3


g 09


a

o c 3 .ft


2


c3 u bo

'ft




i


c3


c5


Dj


c-3




S


M


u





a


0]


CD

03


e


CD

C3




"5


SO


tx


b


> M




j:


>,


>>


>


>>




ft,


Oh



ft



Ph


ft




fi,









IC5



00





ft




CM



fcfl







3







03




fc


N



.3


A



cj^ H


o



<5


<




•&









lO







".


'3



>

S



CD


M



a




M


a



M


o >




>>




a




cj




a







-p




c













in o







P


CD
















u,




>>




ft




O


CI




■b


■b



CS


o

CO




tc




ft




o







ce




CD



as















0)









3










as




m





CD









ft




a


a



















r^



CD




CD


CO ^7 1-1 00


. OS



o ft


03


M S


ft m " 13 m


-PCM

& ft


-p ~

OjCM ^ft


nco

"^CM

co r


1e M ^ft


"as 01

'& ft





-H _£.




r 3


■3* C3«=


"S3"


rft





CD .


CD

35


CD .


CD

<« 


"^^


^« 


<j


—"^3


cj3


CD IS


-J u


CD^£


CD-2


o o_ o t.- 1


gft


Z^


O 3


OCh


OCL


n<;


Ch


Ch


Ch


Ch


•^ W5
















r-»o







p^H




HJ -+J


o


O



O


o


'S'S


r^


r 1



r^


r^


CD CD




ft. ft.




3


"o


"o "



"5


o


CD CD


o


o


o H


>*




^



X


X




XI









cS


c3




03



3


3




3




O*




0"



H


H




a




■b


"bo>






«•


CO CO










CD







a„







O"*







CO CO


■b


■b


bo



b


CD B



C3


coco



o


CO





CO


■b-J







CO








■b


■b


bo


b


b




OS


CO CO


CO


o



CO





CO




bl


tx m £


to





ft


ago






CQ


co O co

CO CO CO


co





CO


CM CM CM


CM







03











CD





















O


ej


.2







C










-p







o


a


3


_C3





kt


>>


>>



CD


_o


ft


CD



'a o


ft



ej


C3


C3


u








C3 CD


fe


a


£


'a


CD


C



c3


cj


09



CD






.3


43


o


_o


O


O








'a


c


3


"3



ja


a


<:



<!



O


ft



Ch


03



CHROMOSOME NUMBERS IN THE METAZOA


49


a


'5



03


00


JOi


111


d



3


•3*



OS 10




£


<j






A


U-C



Oft,


<


ftn


in





H


o


s



©





m






V


b


b


O


o



CO


b


b


C5


o


ce





c


a



o


>


A


03


a


a


>>




03



3


a










an



h



i^


a



a


CO


M a


a ^


a


a


M


a


r- a




r


CM










CO _




eo




S=o



3 3




a


CO



a


00


a

io" >*


OO


a


CDiO


CO ®


o


O0


CDU3 CD 7


.00

a r



CD M


o

60

H



CD £»■

5 -S


2

so


CD M to


>


■a

s

S3


O CD



3



3




9.C


43


3



J3


- 3







a g


a .9


(h



a


M



a <


c3



03 2



£§


s


"c3


.S:0 fc(-H


__;


j:-h



-a


J3


si o



3 CD


ej


,13


CD


o


-1A


O

o


1


"3 o



u


£«>


2

/I —


o —

^h'^


>


J. ~


O t"


N


N-<


<


o


<J o



■3


o



N-fl


■*t<



•^lO







lO
























I -1




-r


















n"




tf*




N






N N















f%


'?'?



>>

CD


£ CD



■|


>>



'S'S


CD


t;



CD




CD CD



c


a


13


o


CD


c S



C


o



c c


u



F*


K


T3


13


W £



M


CD



u l*



0)


a>


























b





b





b



b


b




O





00







OO


OO




W





CM





CM




C-l








o

OO













b











b


b



33



^CM


b


b b



b


b



o


Cft



1




Cft


Cft t-~




OO






OO



B


Cs



CM CM



CM




r^




cm











CM






b-5















CO















CM















M















a















CO















OO















CM













d



ej













o



j3



9














13



13





3



3






3



o



o





J3



J5


~s





ja



J3



-a








a





"o



a s



a

3





o



o o


O

a



'S


CS



o



O


4



a








a £



_2



-2




,0


3



3


3



3









J3



A


-C




(3

O

u

A

o



3

a



13 a


CS



es


.5



a


|S



« 





"3 Q


p.



j

s


S

m



CD

s

w


1

DC




13 C ■J.




3 Oh

-*j

CO CO C co

a

S '3s

CD 1- !

So

csco

o



CO

p

a

CD

CD



c5 3


U

E o

H



CM


b

o

CO



b

o

CO





03

a

03

O >

03 u

CD

O 09

'o

ca

03

o


JOURNAL OF MORPHOLOGY, VOL. 28, NO. 1


50


ETHEL BROWNE HARVEY


oi -3

a a

43 ^







,fl


a



PL|


^00


H



aj





&






■* £r


K

w

K


CD

- 3



C «3


1A





O


tsa-sj


« 



■^ »o




H




S*







w




m



£ £




Oi V




s c









CO


WW


m




M




« 




<!


!*



3


X



K


"3

3 C

W



o







H




H




H




|H




O 1



5


Q




2




C-l




w




H




H

b

o

CO



H










o




2




Q H O









j « 5

o





Q




2




■<





a>


E






w



X>


o




p.


c3

C3





3




O







>>



3




w


Oh




a -*■*




' £



OO



S CO .



M -. o



3 O






z PQJ





<! O,



lO






CO

o







s g



£ 1



a> 3



W 3







b


d




o













a




C3




a.




<N



b

00 <M




b







CM



bD



& M




a



u





















3



jr


3







d



r


o












c


a



r.


a



a.


On




^


a


a



>o


-*


c c



a




4. 9


3

CmJ






3cM .




CO

2





3



"-I0O

a m r


"r


a o

co - O


o


o


CM




■< d



«r


bO


M


■g^N


Nl


N


P4


O




"§ r




tH


u


c_J








3— ■




. 3


3


o o to




-°r-.


cu

a

i

z

Hi




o.2



Zool. Anz. Arch. Nat 1


"3


Intern. M

u. Physi Zeits. wis

p. 503 _ Zeits. wisi

p. 685 Zeits. wis

P. 1


U


a

s



.-3 3P-, © .

^ 3



■*"3


iO





CM


CM






""


r


-"


o o

OS C5


cs


Oi


l-l


tM




o




•C


±s


bj il


b5


b5


CP


03




oc



'(



&


3 3



3






a



P


p


a



A


A


as


C3




15



F

H

h


a "9


a


3


3


3




3



a


CD


0)


CS 0)


aj


o


O


c




Cj



WW


W


W W


w


w


P


P




w









b













3













■-Of













>^3













«■












^ K* 1













>V

























3













•s °













8*













PM






es





c*









a





■*









T3

























'tH













s













CO




b











b


CO


o




b







tn




■«H




>o







es



o

CO

















+




»o







■b





a cs

a

c


3

r


c-i "a!


b






c*




»o II


■2


©







•*




o


-3 CJ


sO










lO













^










cm bo




bD










to O O




o





bD






O 00 OO




o o










CO CM CM




"o





o






a c3 cj




rt co





IQ






OU U




o









o
















3













1


]


o


cs'

cj




si








»



a





o








a



H >>


a




1





s


e



E


G




h





ci





X




JO





G









on







o



.2


'C




•E





'C





"o





CO





o



«,



O


5





s






£



CHROMOSOME NUMBERS IN THE METAZOA


51


A



t-T




rr



r


o







p,


a

<




in


a


c






d






3



fe





c*^ c







>


o


t.-* 1


o



0

fa a




«i


1-1




S





-a a


lO








o





a>"


>>


5

c



|1


o a

u

03

o


T





oa


t




o





n





u








c





a r


00
















|Z(




X





b




■«d









p








b



V,









J,


"b



■b


■^








00



X















a.2













"c







c


















>





!<


c




>>













a


a








03


<




C




a





10



d




o




^ H









B


3




pq



c




o



1

« 









OC




o e

C3







a



L.


iJ




o




,2 a




o -FH CO




s a




ago







-'tf-tS







a ja

J= — -O.




o








OJ







a




o




J5













H






-a



b

O

CO



■b



00


■b

o



>j











o




o




o







"







a







C3

a






E


a




"3










cj




,a




- a




>>



r


$



a.


a,


c <



a a a a ,; .


en oo





vz














a





OP3




•C





c.



2


£


a




+.












=


c


c


c




£


a


i












c


c


c




a a a a



r












X


X


X


X




b b b b






o





















■b






CC


■b 'b -b "b






o
















fafl bfl bD bi) bO



a a a a a









C3


<N


<N


o


CAJ


O







a>






C3

a














03 o3







TS'a « 






a


a m-5








B 1


I


c c



T3

a a


3fe:


c 1


1 s




2 2


o 3 c




£





C




O <


<


<


<


<


-j:


52


ETHEL BROWNE HARVEY


3 a ■ d

m


■9 "* *9


— t.".


= o


a « 


3 c3


n


j= -=




fl C .5 3


73 rt g* en m ,. , O 3 co-c

O.g 5-S Mrr, — p. S 2 0.2 3 m a gw

~ -8 <" ^ ei


X, X X


•§"85

a s m

5 3fl


X X


"b b b b


'b b


egg " a h 9 S H £ 5 a


J3 x. A


-= o


a£ 3 -°33


as u o


-3 a

o


CHROMOSOME NUMBERS IN THE METAZOA


53


a


a


a a


a




h., 25, p.

'omp. Zool. 53, p. 57


a a




"^




CJ


._"


cj


_


CD


. a ■o a N 5

r O

,c


co* >o

■o

> C3

15 s


a S us io " com c-i cni - . cm

£ a

-3 -C " CTJ5




CNI CM

-C J=



o .

en a



." —

Ǥgg


. »o

.-CNI

a •


Q X

■1


a a"

b


Oh

. a


a j.

s a


a a -c .& m t. a • u

a * g ,ga si S| S| -3 s

O O O >2 O




a u .

- - —

O CO^

a =s


a a o o

a s



en


'c a t>

cj C t, o


pa


a

CO


H



j^CN!

ca g

t- o


gS3




as

fc.'— 1 — <" °s


o o 10



c

— ' o PL,


cj— ' e 3

Spq


h'B

c -j q


CS-- '

2 3



a

Q


— • o

3 J

Ph




"*t* -^ ■<*. IOTJ*




■*


Tf -^


















■. i—i






































CO






on










oo











o





bD O



bO M bfl bo bi




M O


Ul tjj







CO



CO



CO


C "



c c e a a




C


C C









a





3 tr"



3 3 3 3 3





3 3





























a




44



ID


^ £



u o o oo




° §


O O




1



s



a c3



o o o o o




O C3


O CD



1



o


a



o


£



s


s


p


s s a as




a o


a s



~



S?


S



!3


s



a







C 6

C3 g


■a c c- 03



















+a



CD









-M











3 U (h


tf


o

a





OT









SQ





e c



CSC m°


a


s s


a a





c




c

















>




!x







-■ -


tu a


> ^ i5


p.


o


qj CD


_CD CD




_0)



CD





o o



a a a is



"° "2


3 3





"3




3





a a



CD




a a


a a





a




a





-2 °



o o o "S-g



o q


o O





o




o







K XI X X
















M M





X X


X


X





X





X






b CN!










b
















1-1










CM































































b b



b b b




b b


« ^



b


b


b


b





cs


CN|



Cs


CN


C\





<M


CM







cs































































CN?











































£ '



b




"b


b





b




b








CN


CN


c



b




CN


CM



<M





o







































CO

























CNI



















bj) bC



bO bO M




to m a


bfi bD



bfl


M


to


to


tc



to


a a



a a




a ao


a a



a


a


a


a


a



a












CO o















CV

CO



« 


o

ec


bO




C*

m« 


« 








« 



CO


CO




CS


CM



CN


CN


c


a

CNI




CN


CM CM



CM



c



o



c




Ol










Vc^









i


















"S 3

g O to




a

O


3

"3





CD

^a








3


5



._;



cj






-m'S'-S




o





"m






rf



C3


d



'3



P






c3 cD-r"




2




s











o



"3






n a '~




CD O


CD





bl

a






"B



'>


■?



S



cj CD






£ * K




a


3





cj




R



D



la


'2



-a


I


c3



c c


,


£


'5 S '£

CD CD CD




CO


m


^z


§


3




S



!



_3


1



_3


'5


'53



c


»


1


O O O




3


3



a


a




"c


,


c



ft


a



"a



j




-s


h h E







o


c




c



c



o


o



o


c


o



c



O CD CD

aa a




'ac


1


C


1


a




r



D



c


a

_2



c





c


£





a c.


Q*











~








c


« 


CD CD CD




.s**


•S?


a


CD


CD


C





a



"3


a


"aJ


C


Q



B


B


S


MhHM HH MH HH




a


w


?


s



~




1



%



§


s



a


54


ETHEL BROWNE HARVEY





CO r

"*


...



.

a ^h


ft


p



a



ft



OS

OS


O -CM

CO ft


OS


02


to


°3 ^


.



'O






a o


B CO


2 >


ft


• OS

.£co "b .


•OS > OS

'a™


. ft

oca

Prfco


d

C3S^3


CI


00 10

CNI

ft

CO pJC


10



"0


a

« 

H

a H

« 


'3 d


O .


2 3&h

^g_ r

c 3N



f a


t> ft



. ft


c



a r



ft


to r

03 —



B


CO

03—'


CO

03j2


o3 £ B


S c

3g


O

g


"a

N


O g


N



O g



B 3

M


_j c3

n


<


1« 


a 3



"a 3§


3 O i-s



3 O


<




K









ir:-^


-r


OS


-*




-*


a















>







CM











00

p


■^ »o


00












Pi.

a to


OS OS


OS


"


00


M b2


b


t,


b» _r



ii


H



si

« 


_g


_s"


r


B B 3 3



CD f


B

3




B


o


§


•p






"5


"K


CD


55


c


^3

CJ


C


P



O



o


o3







CD


CD O



S


c


Fh






iz;


Q


i*£


£


&


fc


s


gg


g


pq


g




g




m j.






M„ o«2











73 a)






^ to %-~*> O

5 3^5 ft.S O

£j= 2 -Ws 2

^§£^ft5







03


2


ts


03 OJ CB



2


-*j



1



2


2



3



_B


.5


T3 c3 B fe 03 g

£ s ™

g Q

-S" S



_B CD

"o


_B

"o


X


"3



a

c


_B

CD

"o



"0


& o


0.

o



ft O


ft



p, a ft 0. 00



ft




X


X



X


« 



X


X


y


X


X



X




b




b






b





c




"b








c<






H



a


CO



S











a




J5













E*
















tH


b


b


2



b


b


b



'■


b '



b



CI





d



CM




c


<M







■fH


ll



»" H


1—1


^ H








•"*


o


^


^7


— '



^


,_7


^T








^


fc


~*


T-H


b



rH


^^


^








^

CM




CO




























a




3












[M




j*













o


b


b


CM



b


b



11


b


b r



b





II



CM


CM









(M



H




-<

'—'



'b







,— '


H

to




b













6
















2 o « o

2 a s


bO


M


mE



bt


bO


bO


CM


bO U bl bO 9



bO


O H S


a


.ft


0.



ft O


ft


ft


II


ft ft ft ft



G


n. <! g


CO


CO


to CO



m c^

co


co


bC


CO


1/


a

too

010



CO


S P< w






C^ CM


CM


CM


ft


e


Cs


CS


— tM



m


g









CM








<!






























Lt




CO


.3

B

<a


CO

3 03 O


.a : 7 3

3 "

H

£



3 Li U

3

E



■3

03


03


03

03





3

(H O

E


B 3

CD




a o


■3


E

CD



CD




03 ft



03 ft


">


oc




03

s


to




a

Cx

to



3


toO

,2 .,



CO

3



3


3


'2


E

bi


D ,.



3 Ml


CD . 03




















&


a


aJL



ft



ft


ft



c



T


c3


SIS


«3



o


o




O



O


O




c



-B



>,



B


c


B B



£ 3


B


B


S


c


c


ft



X



_C3


ji


i3 3



_£3


_rt



T


I


s


CS CJ




"3


"3


O tH



CD U


'cD








03




g


g


g



«5



<*5


s


s


S


c


c


Cm


Ch


&■



CHROMOSOME NUMBERS IN THE METAZOA


55


a


P,


a


.

a


d.


a


a



2,


£



a



6


sS


ra


■^

















o-- 1


rz:


IC


lO


»o


03


'-"


m"


10


ub


■0



».-*


10


to


»o


03 ■

. a


.ffl


10


N"|


<M


CM


CM


O


CM


CM


CM


CM


cm


■*


CM


CM



(M


>> .



a


a °

^


J


J


.i°


,d


J


-3


J


__ r


_CS


_"


_*


6


J3


CM CO


O-MCN

3 n


■*


Sco O »o


ft


a


D


3 p.


a



a


3,


Q




a



a



a „


.


U

o


u o



t>*r



S



O


Eh

O


« 3 c3


O



.


O


3 cj 3


03 > <c


3


cd"°


M


a


£


00 -J


<~


3


§


§


s


",


§


'bo


S


r 1- °

H -a

■ 3 3 =°

co -3h-1

Ph


5


3 §


fa

3 O


►"3


3 b

i-s


- 1 m"3



3 i

^ 1


H3 =i


1-3


10 .•

g


3 y

•-a


-H -CO

to .2 cm

pq


3 O


S 1


— 1

s


3 O •-a


« 


J3

a












CO










-a<


>*


TX



■^


•*




>*



-r


-*



■<*









00


r 1


r*


"


r"


r 1



r^


r 1





05


Oi















p




O


O



hi


bl


tc



bl


bl


bl


bD


bl




bi


ba


2





a


a


3



a


3


3


3


3


C3


a


3



3






3


3


3


3



3


3


3


£


3


3



3



TJ


^3




















O


O


O


3 3


v


O


O


O


O


tH


O


O



O



S


d


>


o








O



C3






CJ



•T


C3


§


S


£


s


p.


a


M


a


§


w


§


s


^


s


M


O



P


+2


4J


































^


fl


3


3


3


3


3


3


3


3



3


3


3


3


3


3



_d



a


_jCB


O


I


<D


01


<D


CD




CJ


O


CD


<D


O






















o


"a


"o


"o


c


O



O


O




c


O


C


O


O




p


5



a



a



a


a



p



a


a


a


a



a


c














q

























-*j






X


X


X


X


M


X


><!


X


X



X


X


X


X


X


X



X














5?




■b



b

















OJ





































00



CO




















'b


b


b


« 


« 


5


b


S


S



b


■b


■b



00


•b


00



■b


CO


"b


1)


b


b

"c


■b


s


"fa


5


^b


"b


■b


c5


%


"b


•b



■b


<M



C*








CM


T








a>

















M^-~




bD bD bfl bfi


bo bD bD hi


b


bl


M bD bd


bD bD


a^



bo


Di Q, P, Pi


a a a a a a


a a a


a a


^J=



a


M


CO





to


en




CQ


CQ










CO



co


CO


CO


o:


CO


CO


CO


CO


CO




CD 8





CM


<m


CM




cq



<M



CI


CM





— a






























^^_^




cd


II



m



3












C3




3




3



CD






3

3 SI) cS

E







§ c3

S.H 4S




3 ci O

O


J3 O

3



"3 3

bD



_a

Fh

3











^s




s



13


3





a

E


I


1


c:


-2

c



E


c

cj e3 c3 cj



c c


3 C


CO

3


3



CO

3



a




c


1- £



c


O O




3


►" c


i< b




j3



c


+■




>




t

M 1

a


c


hj


,3 —


J3


J3





E


_c




c


<L


O


HJ-C


+>




O




5

g


1

>

s

'o

3 » >>


c c a

c


1


« 


1

c


|


u

O O

S3


a


b

c

c


3 3




•0 1


O O

a

CD




H




.c


j:



t~



a


c



'Z


C


t. -m






Xfl


P.


P

a


£


a


a


P.


a


cb aj


tx


V.


CO


CO


w


CO





56


ETHEL BROWNE HARVEY


w o

•z,


o

CO

a



ci

— CO .

tA

00 ft


an Soc. ,

Roy. Soc. 5B, p. 1


a

CI


CO

ft

CO


.r>



co


d


CM

d


ire

CM


d

CM


a



0>


o



2 o


— '


'c .


e .js





jd


a K

a

h

w

K


c

J d



"o3


- Q aH-g


ft c


■as


& ft


Sftft

03S


"3 .pq


C



ft P


a

5



~<m"

>-5


— ' o


O


p ?r

>-3 a


3 3


£ pa

a


S 3

Sffi


PS

"5 pq


a o

>-5


« fa


•re tlire








>o














o















oo


00 IO










-t

>,


o


p ™


ire


■*


^


■*


>














K

H

to

a


CM


CO


CO

•A


»-" CO


bi

a


s

o b£)


c



3 cj

s


b

3 3


c


3





3



O


c


OJ


0) 0) J2 ^


C


C


o



a>



03


G> O


o


o


o


O


a


~



o



s


g


o


§ £


£


s


K


tf « '


w


~


s


§








i.S














Sh cp














o o3 2








m M

PS





c


^2g


"


l^H


c


a


3


3








~^!E








j





_o;


s>


_c



o


OJ


o


o















w



o



"5


o



o



o



p


p


K



a



a


o


ft



c


a


a




o



o


o


> u.ft


o


p


o


p


o


p




■^



+"


■^


"•a



"*a


43


■^




X



X


x


pq


X


X


X


X


X


X









b







Q







b


IM







H


'






o


~ ■







W














H














O


b


b


b

00


b


b


b


b



b


b


b

CM


b



Ci


o>


o



o











«o











a


CO




00






CO





fc














<M














a














H




b











o


b



oo


b


b


b


b


b


b


b


b


'


c;



1


05


^


o


S


t^



5


2


2


H




^*











T.



























o














£














_ a o














2a? o h w

£ 5 *


b6


be



bO


bt


M


bt bt


bB 1



ft



ft


ft ft


S"


a o

en


ft ft ft ft ai 11 'A ai


r~


l^




co


o


CO


CO •"*■


CO


c^



CO








CM





cs


<M


Q O



























la














-* 1 .
















2

o3


"5 !2





£



3 O

o






o

.5*





3




3







'S


3

C3


S3




o






to IB


o



5

ft



■g



o


S

3


=


sj


a> x






o


&










bi








B




3


3 t


3



3


C3 3 .S.S 13 ~ ~ Si W'


x c


"E


^




p.


u




>- c




C3



•r;—


c





-~



-C


3 "5


1



S


2S



_c





o o m



P

-a p


o.s

-S2.P o .


O

o



"3


"3



c


"3






e



^


^2


So u as K


03 —


c


c




S's



|


>



>>


-g<;<ar

a


f



>>



co



CO


co


X


r


CO


CO



H





f



CHROMOSOME NUMBERS IN THE METAZOA


57




a


a






7.


d


~

00


"3


£


.-3


>






£


>


<


03

>


X


oi


« 


fc-.


8


u*


£


.J" CO


01




OcO


g



S



PQ PQ




~h


o



xi



TO


OS


^ H


a




"* X



3


00


OO






CO C


r


Ph


■*


Tf


»o


i>r


Tf



a-*


"3



a


a


a


.


a



t ~ l 03


c <


S^


CO


00

■*


00

■*


6 o

GO


CO



C OJ


,3

1


a

.£co~ MM


© (S3


GO


CO


o




'o'o


2ft


03 fi


J3 ' H


>-i


1-5


gs


J3 c



NN


<


o


<


a


a


Ph


<







©"


,


,








o


o


<





^




5


£





£ t*



p


cr.

p


03


1

03


3 Sic


Ci

p



0> o>





|H



O ©"O




a c


5



to



OJ


OJ





>


u


5 o


a 3*2


H



WW


03


111


o


' o


pi o a


O



£


GO


s


3


s


s




■+J










»2


~*


~« 





— i




_H


13


a





c













O


0)


»





(B













~5


"o


o





O




a


a


a





a




o


o


o





o













X


X


X





X


03



b







■b


-r.









t^



























tO










pq










43











£


b


b





b










r^.




~ H


1-1


1-1


b




1—1







CO




CD


































a















%


b


b

cm



s

©




b







b

CD







bt bjO


M o


bC bfi


be




b« bt




a o


am


n 2


a




a S




to O


en


ffl O





co O




TO-*


to<>


TO-*


CM




TO^J"




CM CM


CMC CO

cm


CM CM


TO




CO CO









©

.2


03 3




ci a







03



03




03







o



"C


o




o













03

'8

03


03

©

"8

03


'8 8



T3 03 8


s

03




8

03



£

f-l

a>

be


g

to M


Ml

03 c


'


03 a

03


03 ©

a




03

a






a


oJ




_03



03


03



O









© 03


Q




[a






3

©







03


c3


_=3pq


3




01



ffl



2



PQ



>-l


Ph




Ph





J.



O CO



a







CM -*






OJ


Q


CM « 





"-"


X


^


r -'




CO



a


a



a


-


"3



»o


„•■



S


CO TO



o



= r



£"5

So

o

o

3


N 00



a



« 


"3


"3


^cS



W



_•


O


+j 3


O


^-'



Jour.

227



o

3


03


in


►J


8g














OO







OC

p


IO


.



CO CO



o



v:


c



o o







be


o








m








O

a


o s

C


4>

© © 'S'S



c to >



03 PS


03 O


GQ


bo


is is

NtS!



4? 00



l





ti


a




O





o


a







>>


o




+J





u


c




4) ffl





o —



^C



•3 CB

a





fOTO


14

o



«" o





O



o



'i©





Cj 03 .


■*


"o



a s





8



a



— . o





o



o



03 C





u


Is




3J=





o-c


>H



CT « 





C U




w





O



X







b




S3


b





■* i



b







CM




3









o









sd









t








o
























ci








03 « 

O



b



b


b


b ■*



~ 0°



d



1


^i




03"







CM



§73

FU.2 C

XI M 03 CM




b




b





b


-r


b


b

CM


■*

T



b




O




CM









be





bt bd £




be


a

en



M



9-S °




a


CO



a



m O gq





a



GO






-!•


Fh



■*



CM <M CM




CM


O ■*



CM



03


03


.3


.S


.3



03










£




'^




u



03


03


03


03



03












3


3


3


3



3




CJ


^OJ


CJ


o



©











£





'C





3


3


3


3



3




o3


03


03


03



03



2


03


03


03


03



03










_03


3


3


3


3



3



"3


o


o


w


a



o



!d


<a



IG



en




Ph


IH


u


tH



U!



a


O


O


o


o



o



<


fe


fe


Ph


tH



Ph


58


ETHEL BROWNE HARVEY



©-H 05

CO ft .'"


p.


r


a




ft


a


|IjngH

a a^tn


o


©


h

a a


M


3



3^


Si

1-5


O

03



p., ca


h-1


a





a





>





3


CO CO iC


3


p





»


  • r

a>



o


^ ^^


fi




O >




O Q) ©



a



-. -

£


bo



6

S


c c 2

3 03 c




MT3


■ ■s >>



M


-a C On


to « eS

z 5



65


o


o a ~



<


© a



S



•a^ o j,



a 65






.= o

03 m


gN*




Ph


IB





b



a



■*







1



o



a





Eh



b



O


b


■*











»





cm





a





>


b



o


•fa


H*


b



(M


1


O






En





10









6





_ a o

2 X h


eX


a



O H H ~ K Z


a


r~




CM



5: •< a

a





cm


>*






<












e~











C3










u






Jl


a




0}

a


~5


"3 t ©


.2


a


3


a


.2*



cj


03


"C







oj


_cj


cj







3

"3


a



©


©


."2



ce


ca






13



o


o


=?



fe


fe





t-~


eg

Eh




o


CO*"


a


CM


c







A


tc





S o




TJ1


■*



1^













«#















©

— '


^J


_jj


CM CM


^2


o






^ H


"""o


^ H



ft . 03


o


cT


C3 <


sj

a


"e3


o

s


ftOS





c



a



n HO Cj £ h

oftu


"3


H

£ CM


a°o

CO


"3 o


d

o

<-H


o N


a





00

m


1-1 C0 M


00

"3 PQ



a * .©Ph


a

<


o


■a .



Cm Oco



1


a ©





"c


S-3 rS

©oft


"o



CO



<


<;


HJJ


s


o


<






(5


§


H



CM













■*



■*















o



p






CM


K3













£




05


OS









t


fcT



©













©


en


©










•&







o


a





CM















M
















cj




03










03



OS



ex




Ph


Ph



>


>






tx -a


a



s

a



a >>


S


E


03

a



o

a






S

3


©

a a


a





03


o


o



'o


'S






03


M


C3



« 



Ph


>


>


m


>


>






m


P3


pq





CD

"3


-3 4

03*0;

>i

~ o -a "a




^ c3


c a


i


a


L

z









a o

X



OJ


a-




oj a

as

=

-r

a

a o


•^


"







M <^


a


"o


a-o





"o


"o


"o





a" os a


T3 « 


ft


oa


IS

F-i


■2


4



ft a


a





I"


I


o


^^


|


a


u


£


G


»H


o


o


o





nT


H



« 







X


X


X

















%


o 03





b




b













CO




r^









- H


■JZ
















O


















>!

































o

















-d





CM
















b




■b







b



b





cm










IC







rH


||




t^







'


rt





,-T











■f



o






b

CO

















































CD

















■«*














b



b


CM



b

crs


b







b


b


b






CO




















t~.-—

















« cj

















>











eX



or ox


bD



ex


IS







CX


ex



a



ao


a



a







a a


a



CO



co O


CO












S3



CO



«■*


CM



r^


ftg







c




<N



CM CM


"- 1



■- 1


a






CN



CM









'


©
















CD

















^^
























3







































o




















_rt










"c3


"C 03



03


03







~




3 ©






©


M



M


tc








CO






N












3







O


a



a


3







|







X!


>



>


>







u


©



si



r3


03



03


03







'53


a:


s



ja



a


0,



_a


J2.







§


© -3


o



"3



"c3


3


"c3


"3



























_o



_o


_o



O


"o







3« 


^3


3


































>>



>.


>>



>>


>.







">>l


">.


>i



Lh




Ih



M








u C


u


Kh



c




6


O



O


O







O


c



c



CHROMOSOME NUMBERS IN THE METAZOA


59


oT


a


^


d



r





a



o rt







cc •




cd





<


"o


fc


— .



>1 .

Offl








u


>i



cc e


00^


s s


J3 P-t


O


pq


-r s

crt


S 9

- o H"5
















ta


■*


■sj


<


N


M




PL,








p


p


_o




CO








0)


a>






-a



-3











3


3


3





O


o


o




§


o







c






^







»^H


X







X













b






P.


a






o








X






X







■b













~*







o








b






b















S £






MMg :






0,0






a


co C


^)0




in








cm cm cm cn












3






3




















S






o


B






s







o


-a






-3








a






3














>,






>>


u







O






a


'Z


a





a lis



cr















■j:


»c





»c



CO CO


oc


oc





»r





CM

10 a


>o



CM

p aa


a



M

a cPh


CO


» a

^-t orT





a





a



£


,-T o


&




o


0 . pq a



■a


"3





"a





Ji




(


ffl 3to o

t-9


s




3 C


lO ^-Ij> •

to MP- a


— '

t-i

<


c


"o

3





<





'o'o o o

S3N


J3 o H











CC







































CM
















■*




















C



c<





,


p


t


lO












bB hn


r




>>

bri -^



i

£






ut






ic:



r


a

3


c




c


1


g


z






~





pp


r -1





a


















c


o





C


T3



a

c . 00


"o









CD CD


^3



c


cd


a '7




5?


£



l





C





- -•

OO


O





7

c a








CI

O


3

o



-*

"2

'S a





"S "2

"o d



z a



C ^ ■ rt j3

an


G cm


4J



CO

"3



"o


"o

a


a

r 13

cs


d

=


3 3 —

3


.2 "3

© o




a _© "o



a o


c

-*

02

CM


a



a o



a o

o -^


1


c


X




a o


3^



X



X





X



X


« 





X





X


X



b






b




b





b




b










03




oo





ec





r—










1 ~ t




"









rt




-3










t-T









o
























b






b



b


b









b












o:


oo









t>









1-1



ci










'"'




ec







_7



oc


tc









CO












w















b







































b


u





b



b b









b




i^



c







oc


oo









t*







02




"-'



OJ


T-H









""






bS




















cr









































M

a













£


3




aa


M



bD bj


bO


all M





M bO


C



M


M bj


o o


a





a a



a





ae




a


o-S





e




»




00





a


o


^


o



CQ O


b<>




i-~




p:


CO









CI



CM



ccoojc


c



CV




CC




UZ


CC





CC



CO oo


CO



CM CO








3



u O

a


c3





oi












C





O

_>

o 3


3

3


03

s

03


.la

S bd


c c




o

a

3

a





3


03



z —



C C




c

c

c


!> 3


Eh Fh

>

3


a

I S

03

F-4


53 S 03 c3

a a

03 03 03 3


s

3

03

3

c:

3




CD >,

J3

a is

a- a v





'>

03


|3 "C

'>

03



oc

C <



3

CO

c




a c


O

Q


o

p


.Sta

P


III







a

c



C Hi




60


ETHEL BROWNE HARVEY




• B


•£


d



'E


a


aft


c;



X


a



•g


■3 a




go


CD


r~


GO


2 co


CO"


3 Ph o

3 ^N


-r

d


CO CO



W O

!5


> 'e


" '(3 .J


d


CM

> oo _


o" 5

. 03

m >

aw


o


'3 .'3 .J



IS

K

a t. w

K


8


— da


13

"3


p ftft as


03 "^

03 C t< O

c


M »a

OJ - X 'bBcoW O



p ap a a





PQ osPQ 3S


O

03


Imp


-j o


£ §S2

03 c ^ ^


.c


l«I«3 S




£


M £


M, £


P9


Ph


U ^


<


« w ^




cm




o — c




ifj C35


3


CM CO ■»*

P P r™



»







O p





OS a tn pq



bi bE el

c c s


- s


bE bS H 3


oc p


CO




bi M bB

3 3 3




J2 2



3 3

u 5


'>



3 3

> >


3

j:

CJ


3 3 3 QUO




ej


o CJ


s


u o


03


(V


CO CO



u




a


s

'cc


a a


Q


a


X.


% % %










-J 3












en O


































o 3 0) .3 t




0)












c


c


c


s


s


rj




3













<


a


<D


Oj


01


OJ


o


"o



OJ



S

K


"o


"o


c


"o


"3


"3


a S5T



"o



a a


a


a


a


a


o °°



a




o



O


o


o


o


-£" s




s

3



X


X


X


X


y.


X


X



X


a


































a o o













b


b


b


b


b





b



Q


t

r



















<


B











Q



Cfi


co



CD


■*
















O












fc












o






















W


a E*











H

tf



b


b


b


b


b


b


b



b






»o


o

<*



t^


<s!




^*


^


'-'


1-1


^


CM



|H




5C


«o


»o


co


"*"


•*


CO



CO*"


~


cm












a




M.S"










b b


„ ° a


b


>b


b




"b




t-~



II a 03


t~


»o


lO

















Eh

m




b"SE









6












z

a H o

2 w s


ho

p. bO


bO


M


be


br


bt


bO

8 »



J K S P- « ft


w


CO


a


a

CO


a

a*


a


a


c


CO

CO



O w o

Q


c^



CO


CO


CM


CM


^


a

c.


CO



z



























03

















-e

3




05


ci


























C3




03


rf












o





S








a








.5


c







o a c

00


c a c e


tf



3

0) c3


R



n


3 -*^

03

a


_H


£





1


'S

c3

.3 a


3


>>

c


o

"3!


"c3 a o

3 0)


c

b


1 1

0.




s


c



6


Cf



■7.


r



a



h


><



£ <d o Ja.a-43

o


bo<


b rt "


bll


CHROMOSOME NUMBERS IN THE METAZOA


61


$




•0


CO



n Ph






ft"







ft "* °*


ft


« a


^


-cS


oa


N co


0)


J2 - O



Js" 1


s


■§•3


3






"S C K


U


O


a 03"


s



■<o


l-l


Ni-3


J




l>- CM





0— 1



QO 00

op


10

)C


.3.3


O


o3 ca


>! SS




s

O


mm


JJ



a a


<D a


Hji-s


> >


GO




CM



















d




•BPj


"0 a



M O

J3.3


c a

03 fr-'"* "c3-5'o







03 »<


O ft


X




X


b





OO





'""'





r^
















OS





I -1


b



O


O0




O



" H



« 


II




03






b

QO







b


OS


b


a


O


II



1





j


00






00






S




t^


Mg





ft



o__


H


<=o








mo


cc re



CM CM


ftCO CO







03













r3






3







>>


T



CD


S3



c


cl


3



3


3















as


OJ


<



X


m



i-3



^_


ft



■*


09 (-^






ft



CO







3




OS


ftoo




CO

•O




•0


5


03


""^T*



X


ss 6


-r


rt


a <

cj



cm


"0

m


_;


< 6


53 co



a


10 ■


■6

a >


ft

Ci


a



<



CO CO


_o



St3




3


_r 3

■3S


r. X


"3



4-*


3

m



i

eo


"a O


&*~.



00


O


m


C

a


m


•o



z



03


ft


s<


03


is?


s:

-"r



"0



O




Oh



J


mm



5


d "


m



p


B



a"








"3
















"0



3 O

m



3


m


^




CO IO




S








00



r"


<!



















oc





>c


a" a"

c3 03







a


S



S


3



"



2


a


p



c3

a '3


-0


■0 y r ns23

s -°



03



>>


C3


as

sj 73



J5


'BX 0

CD

■a

CD



pq


m



ft


m



O


55



3


m


S













co 50

c; a











^




"°.2


s









CO




'>!2


iS










a




^.b


e













T3








X




"3

0.




>>ja

io-^

8a


"3

a



































V,




«!


X










b






s

"2


b













5.









CO



























V



















O















b

00








b

CO





b

r. 00





b b




b








b



5


cm


b



CI


b




b


b



oc



J


cm

CM


■0



CM





00


o>










bC





bt










ft





ft










CO














CO





r^










CO





CO







03



03






03




















a



c


3

"S


ca

■2.


03 -m

E

a


ft


03

03



03




03

. a


"o


C a

c


,



a



I


3


00 CO

2.2


3


— 3


a




cS


03



LE



'2


3


ISIS


IS


CD

ft


CD

a




CD bS


CD

to



c c

a



c



O O CO



O









-a


.c-e 3-a


"3


"o




3


3















C




J


jm


-;


co


cc





CO


X




-*



00







-H fcQ


•O



ft ft


ft






CO CO




^^

-»<









S CD


a



<N


<!









O C3







n«;


SJ






pp


T»<










fO



e a













OO


CO












a







CD










ft




O







X



n










R







B,













b




CO


b




00






CM







b •


b




00







M




ft






ft




■0


^2 O

ft



















cd




>


a






3


3



3


a



>>


CD



-a


>>









Cm



0



62


ETHEL BROWNE HARVEY


< O « 

1 °

Q 8

? 8

P-l (H

O Z

rt o

w « 


H o

Z H K H

W

K


1 ft

-ft

c



(S W >

a

H m

pq O


o p

>,

fc. o

a

O tJ)

fl

O


in M B ■< S H PS



g

1


■b


o

1

Q

Z


■b


o

03


•b


6

z

Q W O

£ n " ° & U

z


J?

CO

hOoooo oo

CM


m

H

3

H

P.

to


D


P

c a


,


<!

"*< - 1

« cj § §

ft o » o

H (h §■,£.

H . .2

O e -< ^

O CS


ft


d


n


a


OS



^




ft


da




CO


cT oo


o


CM


CO CM

ftft


csT



05

o

CM


CM


Scoo io

o

05

o

CM


„ r


_■




^"oo


^ r


a


ft



fcH

aS no >

3 u - OS

3nS


ft


o

o N

1


o o

SI


o o


a

o


CM CM

fc. fc<

,3.3

C3 oi 1-5 1-5


Q

o

N

'1


00

PQ


pq


a u o


3 PQ



"5

~* -^ 00


■+i CO


goo

1-5


"3 "5

o o


.05


"o


t!05


3


N


[SJ


N


NN


N


pq


« 


1-9


PQ


5















p






05




05





■*


lOOS




O


T)<



O


p

s"


oo p


a '53

c go


A


Op

oJ c2

2 2 "3 "3


o:

a 03


OS O

a> W


-0 o o

_^

"3


0)


OS O

'Si fc.


-a" o o j* 3


ti


3


o



o o


C3


g


03


E


a


S


§


m


PQ


pq


^


M


PQ


W


m






■m























o












fi












— c























OJ












a <k












O C3












o ft












3 ci












T3 C












O C3












r^























'be*












toco












'bo











te CO

















^3












O



Ot

CD


o


t>


o


"bo


O

-t"


&



o


fc*

a.




co


s



CO CM


V


CO



T}« 

























CJ










i

o







M




"3









ft




CO




'o






cq




J




■*



ll ^















11 a















a


.2


a

3






cj


e!







c3 o





















c3


03







o^


3


O






a


a






e


S3^


cJ


s

0)





3


IS


A


_d



o—« 










^


S'O


08

fc. O



03 m






a>


p


s


ft 2


a


01 o^


a



.2


2


.2


.2


E


03 g


,4


_3


2 a s




a


'.5


'■4J


+3


p "3-3


ft


'C


•-3.5->


t


D h


» 'I



.2


.2


>


g|



c

0)

3


C In' C


oi



03 H

13



03

3

a

0.


S3

a a o


c


.2W


O O


>>>>>*


>


^ >


i >>


>.


1 C


o


H


ffiH


w


n


n


w


n


PL.



a



CHROMOSOME NUMBERS IN THE METAZOA


63


M Tj*









CO








S n



m












o


a


o


® m



SSJ





3 o



CO


-wN




c _•



£









%*>



•S"»


N



N





en














o



3







u


ffl



pq


o










o














o


o




■*












u




00




ON







q











>











,fl






a






0>






a




«*


C3


a


e




3


</




a




3





,Q.S



3


4^


p



H




o" a


a






!>■


oj M



P.



<



O


.


a


N


& s


ui


£


a


1




. CO


„■* goo

< 1-9


■g<M



CO















^ '5b





oj a

3


w w


5s


o



CO




o


o



o


00








L


o



©


o







03





o




CS


3


mr i



3


G CD

O


es


en



|


05

s


r


3


I


s~





ft

• a


o



a


a


03



<!


O


O




aa


a


o .





03





3 b


w -co o "^ o •

N a


as


3Ncm


a


PQ




&



o





&



o



a


o


03



.isl


&



o





M




CO


pq


"b


CHOI


+


"- 1


-*


,0





to



a



■b



CI



■"■•



« 







t


a


■b


CM


+o


1-1



o


tin




a



m a



M O O



ao <n



t0 -*„-*





"•"§«•§





a a







.c





C3



X!





3


o


a


-Q



O


3


a


a





a


d





a


a


EXPERIMENTAL CONTROL AND MODIFICATION OF LARVAL DEVELOPMENT IN THE SEA URCHIN IN RELATION TO THE AXIAL GRADIENTS

C. M. CHILD Hull Zoological Laboratory, University of Chicago

EIGHT PLATES


The literature of experimental teratogeny in embryonic and larval development is extensive and a great variety of external factors has been employed in experimentation along this line. While the results have often been of great interest, no general basis for interpretation has been reached, particularly as regards the relation between teratological form and the action of experimental factors to which the developing organism is subjected as a whole, and not by local application. It is a familiar fact that even such factors affect different regions or parts difr ferently or to a different degree, and while such differences indicate the existence of local differences of some sort, there is, in general, a very evident lack of specificity in the action of different external factors on the course of organic development. Similar teratological forms may often be produced by many different agents and conditions.

The present paper constitutes one step in the attempt to correlate certain types of teratological development with dynamic conditions which are characteristic and fundamental features of the normal organism. It establishes a basis for the control, modification and prediction of the course of development in the sea urchin, and the facts already at hand indicate very clearly that certain types of teratological development, as well as certain characteristic features of normal development, can readify be interpreted on the same basis. More specifically this paper is a demonstration of the effectiveness of the axial metabolic gradients as dynamic factors in the development of the sea

65

JOURNAL OF MORPHOLOGY, VOL. 28, NO. 1


66 C. M. CHILD

urchin. It shows how, and to what extent, it is possible to control and modify development by means of the differential action of external factors on different regions of these gradients.

THE NATURE AND PURPOSE OF THE EXPERIMENTS

The existence of an axial metabolic gradient, at least in the major or polar axis with the apical region as the region of highest rate of reaction, or, perhaps, as possessing the capacity for the highest rate of reaction has been demonstrated in many species of animals and plants (Child, '13 b, '14 a, '15 a, '15 c, Chap. Ill, '16 a, '16 c; Hyman, '16), including the developmental stages of the sea urchin. This demonstration has been made possible by the differences in susceptibility to various agents which are correlated with the differences in metabolic rate and protoplasmic condition at different levels of the gradient.

So far as this relation between susceptibility to inhibiting agents and metabolic rate has been investigated, it is as follows. In concentrations of cyanides, narcotics so far as tested, acids, and, under certain conditions, alkalis, which kill, not immediately but rapidly enough so that the organism does not become acclimated or acquire a tolerance to them, the susceptibility varies directly with the general metabolic rate or with the rate of certain fundamental metabolic reactions. To low concentrations, to which more or less acclimation is possible, the degree and rapidity of acclimation vary in general directly with the metabolic rate — with certain exceptions (Child, '14 b) — and within certain limits of low concentration the parts least acclimated die, while those more fully acclimated remain alive, so that in the long run the susceptibility varies inversely as the metabolic rate (Child, '13 a, '15 b, Chap. III).

In Planaria, for example, it has been shown (Child, '13 b) that to the higher concentrations of cyanides and narcotics the susceptibility decreases from the anterior end posteriorly, while in the very low concentrations the degree of acclimation is greatest at the anterior end (Child, '12), and in the long run susceptibility is greatest at the posterior end and decreases anteriorly.


LARVAL DEVELOPMENT IN THE SEA URCHIN 67

From the metabolic point of view, -acclimation consists in the gradual increase in rate of reaction in the presence of the inhibiting agent, and the inverse relation between capacity for acclimation and metabolic rate means that, the higher the metabolic rate, the greater the capacity for gradual increase of rate after the first inhibiting action.

Recovery after the temporary action of an inhibiting agent consists, like acclimation, in the attainment of a higher metabolic rate, but the rate attained is much higher than in acclimation because the inhibiting agent is completely removed. Where the action of the inhibiting agent has not gone so far that recovery is impossible, the rapidity and degree of recovery like acclimation vary directly with metabolic rate; the higher the metabolic rate the more rapid and complete the recovery.

If these conclusions are correct and if axial gradients are present in organisms and are actual effective factors in development, these relations between susceptibility and metabolic rate afford a basis for the control and modification of development in two opposite directions. First, by the use of concentrations of an inhibiting agent sufficiently high to prevent acclimation to the reagent, or concentrations and periods of action sufficient to prevent recovery after temporary subjection, it should be possible to inhibit the regions of higher rate of reaction in an axial gradient to a greater extent than those of lower rate. If the rate of reaction in the apico-basal axial gradient decreases from the apical region basally, the apical region should be most inhibited, the basal least under such conditions. Second, by the use of low concentrations which permit some degree of acclimation, or by temporary action of concentrations whose effect is readily reversible, it should be possible to obtain a greater degree of acclimation or more rapid and complete recovery in the region of high rate, i.e., the apical region, and so to inhibit its development to a lesser extent than that of the regions of lower rate. In short, it should be possible to produce, in the one case, a gradient or gradients in inhibition of development, and, in the other, gradients in acclimation or recovery. In fact, by determining proper concentrations and times of exposure, it


68 C. M. CHILD

should be possible, not only to produce differential or graded inhibition or acceleration of development along an axis, but to determine the position of the greatest degree of inhibition or acceleration, either at the high or low end, the region of highest or that of lowest rate in the axis. To obtain a particular degree of differential inhibition, the procedure must of course be varied somewhat according to the toxicity of the reagent.

The experiments on the developmental stages of the sea urchin were undertaken with these ideas in mind and with the purpose of demonstrating that the axial gradients are determining and effective factors in embryonic development. The results of the experiments have completely realized expectation and leave no doubt as to the fundamental character of the role played by the axial gradients in the development of the sea urchin. Some similar data on the starfish, which I hope to supplement by further experiments before publication, indicate that the relations are essentially the same there as in the sea urchin.

THE AXIAL RELATIONS IN NORMAL DEVELOPMENT

Throughout the paper the terms 'apical' and 'basal' are used to designate the two ends of the apico-basal, polar, or major axis, the apical end representing the apical or 'animal,' the basal end the 'vegetative' pole of the egg. In speaking of metabolic gradients, the ends or levels of different metabolic rate in a gradient are distinguished as high and low, or higher and lower ends or levels.

According to Boveri ('01 a ,'01 b) the micromeres arise at the basal pole, and the micromere region gives rise to the mesenchyme, but Garbowski ('05) has shown that polarity of cleavage and of the later stages does not always correspond to the axis indicated by the pigment ring in Strongylocentrotus lividus. The elongation of the blastula occurs in the direction of the apicobasal axis, and the apical pole of the blastula represents the apical pole of the egg. Before gastrulation, the basal region of the blastula wall becomes thicker than the apical (fig. 1). In the gastrula (fig. 2) the apex of the conical body represents the apical end of the major axis, and the blastopore region the basal end, while the invaginated entoderm represents a still more


LARVAL DEVELOPMENT IN THE SEA URCHIN 69

basal region of the egg and blastula, and the mesenchyme the most basal region.

The transformation of the gastrula into the pluteus begins with change from the radially symmetrical form of the early gastrula to a bilaterally symmetrical form. The basal outline of the gastrula becomes somewhat triangular instead of circular (fig. 3 A, basal aspect) and the apical region appears to shift toward one end of the longitudinal axis and becomes the oral lobe (fig. 3 B, lateral aspect). Meanwhile the apical end of the enteron unites with the body-wall at the point near the apical pole where the mouth develops, and the enteron is marked off into three regions, the oesophagus, the stomach-intestine and the rectum or anal region. Also during this period the skeletal rods appear, the body elongates and the anal arms begin to develop (fig 4, lateral aspect). Further development consists in the continued elongation of oral lobe, the appearance of the short oral arms and the elongation of the anal arms and of the body. The fully developed pluteus (fig. 5 A basal and fig. 5 B, lateral aspect) possesses a very definite antero-posterior axis, the anterior end being the broad end bearing oral lobe and arms, with the ciliated band running over the arms and oral lobe and the body margins between them, the posterior end, the opposite tapering end. Since the oral lobe represents the apical, and the anal surface the basal end of the original apico-basal axis, it is evident that this axis is no longer straight but curved, in consequence of the shifting of the apical region toward the anterior end. The course of the alimentary tract is in some degree a record of this change. In the analysis of the differential inhibitions it is essential to keep in mind this modification of the original apico-basal axial relations, as well as the antero-posterior relations of later stages.

METHODS OF EXPERIMENT

The reagents used in this series of experiments were potassium cyanide, ethyl alcohol, hydrochloric and acetic acids, sodium hydrate and ammonium hydrate in various concentrations. The eggs or embryos were placed in the solutions in


70 C. M. CHILD

500 cc. or 1 1. corked Erlenmeyer flasks filled almost to the cork, and solutions were renewed every twelve hours where the period of action was longer than that. It has been previously determined that development would proceed normally in such corked flasks in well aerated water, renewed every twelve hours, provided the number of eggs or embryos was not too great, and particular care was taken in the experiments that the number should not be excessive. In the experiments involving recovery after temporary action of the reagent, lots of eggs or embryos were returned to sea water at certain intervals, with four or five changes to remove as far as possible all traces of the reagent.

THE EXPERIMENTAL MODIFICATIONS OF DEVELOPMENT

If the above statements concerning the relations between direct susceptibility, acclimation and recovery and metabolic rate, are correct, and if metabolic gradients are actual and effective factors in embryonic development, it is evident that three possibilities exist for the modification of development by the action of inhibiting agents. These possibilities are: first, direct differential inhibition of development by relatively high concentrations or highly toxic agents, in which case the degree of inhibition should vary directly with metabolic rate; second, indirect differential inhibition through differential acclimation to lower concentrations or less toxic agents, in which case the degree of inhibition should vary inversely as the metabolic rate; third, indirect differential inhibition through differential recovery after temporary action of the agent.

The experimental results realize expectation. In the forms produced by direct differential inhibition, the development of apical, anterior and median regions is more inhibited than that of basal, posterior and lateral regions, while, in the forms resulting from differential acclimation, the development of basal, posterior and lateral regions is more inhibited than that of apical, anterior and median. The forms resulting from differential recovery are, in general, similar to those resulting from differential acclimation.


LARVAL DEVELOPMENT IN THE SEA URCHIN 71

In short, these experimental methods produce two opposed types of teratological forms with various gradations between them, and the production of the two types can be controlled to a very considerable degree, the differences in susceptibility in different individual eggs and different lots being the chief limiting factors in control. For convenience the forms produced are described under the four heads: direct differential inhibition, differential acclimation, differential recovery; differential inhibition with general recovery; and consideration of the question of control follows the description. The figures of the teratological forms are semidiagrammatic and are intended to show the form, axial relations, and chief structural features, details of mesenchymal distribution and slight skeletal variations being usually omitted. Wherever the distinction has seemed necessary or desirable, the middle region of the enteron has been drawn with a double contour, as its walls are thicker than those of the oesophageal and rectal regions which are indicated by single contours. All figures were drawn directly from living material.

THE FORMS RESULTING FROM DIRECT DIFFERENTIAL INHIBITION

The results of direct differential inhibition are most clearly seen where the development takes place in KCN, for little or no acclimation to KCN occurs within the short period of development from egg to larva. In NH 4 OH, development also occurs without appreciable acclimation. Some degree of acclimation to NaOH occurs during development, but in alcohol and acids acclimation occurs so much more rapidly and completely than in the other agents used, that concentrations high enough to produce a large percentage of partial or total death must be employed to produce the forms characteristic of differential inhibition.

Considering in order the various degrees of departure from the normal form, the first appreciable differential inhibition appears as a slight change in the proportion of parts in the pluteus. A comparison of figures 6 A and 6 B, drawn from a very slightly inhibited pluteus, with figures 5 A and 5 B, (a


72 C. M. CHILD

'normal form) shows that in the former the oral lobe is shorter and less developed, the anal arms slightly shorter and the angles of divergence between the lateral skeletal rods and between oral lobe and arms are slightly smaller than in the latter, while the length of the body from the base of the anal arm is essentially the same. In these plutei the development of the oral lobe, which represents the apical region, is most inhibited and the anterior end is smaller in both dimensions than in the normal animal, while the posterior region is fully developed. Forms such as this, in which the angles of divergence of the skeletal rods are less than the normal, are called, for convenience, narrow-angled forms. These differences may appear in different degree in different individuals of the same lot, in some the only difference from the normal being a slightly shorter oral lobe, while in others, where the oral lobe is inhibited to a greater degree, the angles of divergence, which are indices of the development of the anterior end in the direction of the minor axes, are also smaller than in the norm. Where the degree of differential inhibition is so slight, the earlier development is merely somewhat retarded without appreciable departure in its course from the norm, and the differential inhibition begins to appear only in the later stages.

A somewhat greater degree of inhibition produces forms which attain the condition of figure 7 A and B, but develop no further. Here the larva remains small, the development of the oral lobe is completely inhibited, the anal arms remain short, and the dimensions of the anterior end, as compared with more posterior regions, are much reduced, so that the angle of divergence between the skeletal rods is also reduced as compared with the normal form. The entoderm, however, still undergoes differentiation into three parts and the mouth forms. It is evident that in such cases the development of the apical region is most inhibited and the development of the anterior end more than that of the posterior end.

Where the degree of inhibition is still greater, the differential inhibiiton may appear even as early as the elongated blastula stage, as a relative decrease in apical and increase in basal di


LARVAL DEVELOPMENT IN THE SEA URCHIN 73

mensions (fig. 8). The gastrula formed from a blastula like figure 8 is natter, i.e., apical inhibition is greater than in the normal form (fig. 9). Occasionally, in the more extreme cases, the entoderm evaginates instead of invaginating, and an exogastrula results, but in such cases, so far as my observations go, at least some of the mesenchyme cells pass into the blastocoel. Figure 10 shows the beginning of exogastrulation, and figure 11, an exogastrula. Under no conditions, however, has exogastrulation been observed in more than one or two per cent of the inhibited forms.

In the more extreme degrees of inhibition, the form of the body becomes more nearly spherical, the entodermal differentiation more completely inhibited and the skeleton more rudimentary or completely absent, although, even when they are entirely unable to form a skeleton, the mesenchyme cells are not dead, but persist in the blastocoel. In most cases where a skeleton forms, the angle of divergence between the skeletal rods is narrow. Figure 12 A and B, shows a form with rudimentary narrow-angled skeleton and figure 13, a form in which the skeletal rods are almost parallel. Figure 14 is an anenteric larva in side view with rudimentary skeleton, a form resulting from an exogastrula; figure 15, an askeletal form with differentiated entoderm, and figure 16, a still more inhibited askeletal form, in which the entoderm never develops beyond the stage of the spherical vesicle attached to the body wall. In forms like figure 16, apico-basal and other axial differences in the ectoderm never appear, and the blastopore usually closes completely, so that it is impossible to determine whether the position of the entodermal vesicle represents the basal region or not, and certain cases of differential recovery to be described below will show that the entoderm may entirely lose connection with the blastopore region. In the case of exogastrulae where the entoderm is entirely lost, the askeletal forms develop into thin walled ectodermal spheres containing scattered mesenchyme cells, and without any visible axial differentiation in any direction.

In these spherical forms all axial differentiation beyond that of the gastrula is inhibited, and it is of interest to note that in


74 C. M. CHILD

these cases the mesenchyme cells are scattered irregularly and do not aggregate in the basal region, as they do to a greater or less extent in less inhibited or normal forms. Evidently the localized differences in the ectoderm which supposedly determine the distribution of the mesenchyme are not present to a sufficient degree to be effective.

The most extreme degrees of differential inhibition are partial forms in which a portion of the body has been killed. In these cases death proceeds from the apical region, and the part which remains alive therefore consists of more or less of the basal region (Child '16 c). When such partial apical death occurs before the gastrula stage, the gastrulae are small, with a disproportionately large entoderm, though not always as large as the normal (figs. 17 and 18), gastrulation itself apparently being inhibited to some degree. When partial apical death occurs after gastrulation, the result is not essentially different. These partial basal forms show no ectodermal differentiations, remain spherical, the blastopore closes, and the entoderm remains as a spherical vesicle (figs. 19, 20, 21). As the figures indicate, they vary in size according as they represent a larger or smaller fraction of the apico-basal axis, and in the smaller forms, such as figures 20 and 21, the entodermal vesicle completely fills the blastocoel, because the whole of the entoderm, and only the basal portion of the ectoderm, remain alive.

In all these spherical forms, whether whole (fig. 15) or partial (figs. 19 to 21), the entoderm often loses its coherent epithelial character after several days of life without further development. In figure 22 this process is indicated. The cells become scattered about the blastocoel, and such forms are distinguishable from the anenteric forms resulting from exogastrulation, only by the larger amount of cellular material in the blastocoel. I am inclined to regard this disintegration of the entoderm as connected with its lack of functional activity, and it may be that obliteration of the metabolic gradient in the entoderm is also concerned. At any rate, this entodermal disintegration has been observed only in the inhibited forms, where the entoderm is cut off from the basal region and does not differentiate.


LARVAL DEVELOPMENT IN THE SEA URCHIN 75

The forms shown in figures 6, 7, 12 to 16, and 19 to 21, are the final stages of development attained in the different degrees of inhibition. This does not mean that the animals die at these stages, but simply that no further development occurs, although life and more or less movement may continue for days. It is evident that the various degrees of inhibition are likewise progressive steps in the elimination of the axial relations as effective factors in development and differentiation. If the axes are fundamentally metabolic gradients this is easy to understand, for, since susceptibility varies directly with metabolic rate, the decrease in rate is greater in regions of higher than in regions of lower rate, and the metabolic gradient is therefore more or less completely obliterated by leveling down according to the degree of inhibition. When this obliteration by leveling down proceeds to a certain point, the metabolic gradient ceases to be an effective factor in development and differentiation, and the animal becomes physiologically anaxiate, and, therefore, development and localized differentiation cease, although life may continue for a long time.

THE FORMS RESULTING FROM DIFFERENTIAL ACCLIMATION

Differential acclimation to an inhibiting agent is of course preceded by a greater or less degree of differential inhibition which, however, is always less than where the action of the agent is sufficient to prevent acclimation. The effect of differential acclimation on the course of development and on the form of the body, differs in degree, according to the degree of inhibiting action and the rapidity and degree of acclimation, but the characteristic feature of the morphological modifications resulting from differential acclimation is that they are opposite in direction to those resulting from differential inhibition. The regions most inhibited in the one case show the greatest relative acceleration and over-development in the other.

Of course where acclimation is slight in degree or occurs slowly, differential acclimation may only partially obliterate the differential effects of inhibition, so that the resulting form merely shows a less extreme degree of differential inhibition than it


76 C. M. CHILD

would if no differential acclimation had occurred. In such cases the form is not an adequate criterion, though increase in motor activity and rate of development may indicate that some degree of acclimation is taking place. But when the axial relations of differential development are the opposite of those resulting from differential inhibition, the form is sharply distinguishable from the form produced by differential inhibition, and it is with such forms that we are at present concerned.

Where the direct inhibiting action of the agent is not very great, differential acclimation does not produce any extreme modifications of form, but appears merely as a change in the proportions and angles of divergence of parts of the plutues. Figure 23 A and B, shows a differentially acclimated pluteus in basal and lateral aspects. A comparison of this figure with figures 5 A, B, which represent the usual form of plutei raised in pure sea water shows that, in the acclimated form, the oral lobe, i.e., the apical region, is longer and broader, particularly at its apical end, the body is shorter and less slender, and the angles of divergence between the two anal arms, between anal arms and oral lobe, and between the short arms on the oral lobe is greater than in the sea water forms. Moreover, a comparison of figure 23 with figures 6 A, B, which show results of slight direct inhibition, makes the opposite character of differential inhibitory and differential acclimatory changes strikingly evident.

Not infrequently in these wide angled forms, the angle between the anal arms is somewhat wider than between the posterior skeletal rods, as in figure 24. This indicates a greater degree of disproportion in development between the anterior region and the more posterior levels than in figure 23, i.e., breadth of the anterior region is increased out of proportion to that of other levels of the antero-posterior axis, and the angle between the arms is therefore wider than that between the more posterior portions of the skeleton.

In KCN and NH 4 OH, where acclimation is relatively slight and occurs so slowly that development is usually completed before the differential effects of acclimation become marked, even the slight changes of form shown in figure 23 do not usually


LARVAL DEVELOPMENT IN THE SEA URCHIN 77

occur, but in alcohol and hydrochloric and acetic acids acclimation occurs much more rapidly and to a much greater degree, so that differential acclimation is often evident in the early gastrula stage. Consequently, with these agents much more extreme modifications of form can be produced through acclimation. NaOH is intermediate between these two groups as regards the forms resulting from differential acclimation.

The earliest marked indications of differential acclimation which have been observed occur in alcohol and acids, and consist in increase in the relative size and growth of the apical region of the gastrula. In acids this is very commonly a tapering prolongation of this region (fig. 25), while in alcohol it is more often rounded (fig. 26 B), but both forms may appear in both agents. Irregularity of outline and roughness of the external surface of the basal region of the gastrula (figs. 26 .A, B), is particularly characteristic of the action of alcohol, though often produced to a lesser degree by the acids. It is evidently due to a decrease in the normal epithelial coherence of the cells in this region, but' as development and acclimation proceed, it almost or quite disappears.

Where the apical region undergoes a relative acceleration in stages as early as the gastrula, disproportion between it and other parts increases as development proceeds. Figures 27 and 28, are side views of prepluteus stages showing this disproportion. As compared with the normal form at this stage (fig. 3 B) the apical and anterior regions are relatively greatly over-developed, and basal and posterior regions, including skeleton and arms are greatly under-developed. These alterations are exactly the opposite of those resulting from differential inhibition (fig. 7 B). In the acids, where acclimation occurs more rapidly and the differential features are more marked than in any other agents used, the most extreme modifications of this sort occur. Figures 29 and 30, A basal, B lateral aspect, show cases from acid in which there is enormous over-development of apical, as compared with basal regions, and in which the anterior region is so much over-developed that the arms approach or attain an angle of 180°, while in more posterior regions the angle of divergence,


78 C. M. CHILD

although wider than normal, is not as great as that between the arms. In these cases the effect of differential acclimation has been extreme in the apical and anterior regions, but relatively less in more basal and posterior parts. Figures 31 to 36 show the final stages of development attained in the more extreme degrees of differential acclimation to acids. Figure 31 A is the basal and figure 31 B the lateral aspect of a larva with large oral lobe, small body and no arms. The body is short and broad and very wide angled. Evidently apical and anterior regions are relatively over-developed, basal and posterior relatively underdeveloped. In figure 32 A (basal) and B (lateral), these modifications are more extreme. The oral lobe is relatively longer, the body shorter and broader, and the angle of divergence wider. Figure 33 shows a still more extreme degree of modification, A being the basal, B the lateral and C the anterior aspect, and figure 34 shows the most extreme modification possible in this direction, A, B, and C being, as before, basal, lateral and anterior aspects. Here the oral lobe is larger than the body, and the skeletal rods are united to form a single straight rod lying transversely, and passing anterior, instead of posterior, to the anus. The angle of divergence between the rods is here 180°. These forms in figures 31 to 34 differ from those in figures 29 and 30 in that the angles of divergence of the skeletal rods are uniform throughout, instead of being wider anteriorly than posteriorly. This means that in figures 29 and 30 the differential effect of acclimation is more extreme anteriorly than posteriorly, while in figures 31 to 34 it has extended far enough posteriorly to determine a uniform angle of divergence of the whole skeleton, instead of a wider angle anteriorly than posteriorly. In figures 35 and 36, A, B, C, forms similar in outline and axial relations, but without skeleton, are shown.

It is evident that all these forms are the result of relative differential accelerations along the axes. Acclimation has made the slope of the axial gradients steeper; i.e., the regions of highest metabolic rate in each gradient have now a relatively much higher rate, and the regions of lowest rate a relatively much lower rate then under normal conditions. Consequently the greatest de


LARVAL DEVELOPMENT IN THE SEA URCHIN 79

gree of over-development, as compared with the norm, is in the regions of highest rate, i.e., apical and anterior, and apparently also median; and the greatest degree of under-development is basal, posterior and apparently lateral. In consequence of these changes in relative metabolic rate along the axes, the larva is transformed from the antero-posteriorly elongated normal form (figure 3 A) into a broad, transversely flattened, form with short antero-posterior axis ; and all stages of this transformation appear.

In cases where the degree of inhibition is somewhat greater, differential acclimation in acid gives rise to forms like figure 37, which shows basal (A), lateral (B), and anterior (C), aspects of a characteristic acid form. In this larva the ciliated band, which, in the normal larva and the less modified forms, extends as a continuous band around the margins of the anterior end and over the oral lobe and basal arms, is differentiated only in the apical and basal regions, as indicated in figure 37 by the shaded bands. The apical portion of the band extends around the rudimentary oral lobe and a short distance basally, while the basal portion extends from the medial anterior region posteriorly more than half way around the body, and the lateral portions, which normally connect these apical and basal portions, are not present. In fact, the basal portion of the ciliated band in such cases forms a more or less complete ring around the basal region. This condition recalls the condition in the more primitive type of echinoderm larvae where several ciliated bands surround the body. Apparently in such cases as this, where the anteroposterior axis and bilaterality are practically obliterated, the local metabolic conditions determine that the ciliated band shall develop around the basal region, instead of over the arms and around the lateral margins of the anterior region. This condition also appears frequently in recovery forms (pp. 85, 87, 88).

When the direct inhibiting action of the agent is somewhat more lasting, and acclimation occurs somewhat more slowly, as in alcohol, the changes resulting from differential acclimation are, so to speak, superimposed on the changes resulting from differential inhibition. In such cases, therefore, various combina


80 CM. CHILD

tions of differential inhibition and differential acclimation appear. The form shown in figure 38, for example, is, in its basal aspect (A); a characteristic case of differential inhibition, resembling figure 12 A, with the inhibition decreasing from the anterior to the posterior end, so that the body is relatively narrower and the angle of divergence of the skeletal rods less than in the normal (fig. 3 A).

The side view of this larva, fig. 38 B, is very, different from that of the differential inhibition, figure 12 B. The oral lobe is large, with a rounded knob on its apical end, and the ciliated band develops only in this apical region. Great elongation in the apico-basal axis has occurred, and the enteron is attached at the mouth region, but not at the anal, and the rectal portion is absent.

Figure 39 is a side view of a form in which differential acclimation follows a somewhat greater degree of inhibition and consists chiefly in the apical outgrowth and the broadening of the anterior end.

The observation of many similar cases has shown that the course of development in cases of this sort is as follows: during the earlier stages of development the action of the agent is directly inhibitory, with the usual differential effect, but, at about the gastrula stage, differential acclimation begins to be apparent as a relative acceleration of development in the apical region, as in figures 25 and 26 B. From this apical region the relative acceleration progresses basally, but is more marked in anterior than in posterior, and apparently also more in median than in lateral regions. The basal region undergoes acclimation so much less rapidly and less completely than the apical that it shows no secondary changes before development comes to a standstill in the solution.

When the apical elongation begins the entoderm remains attached to the region where the mouth is to form, and it may break its connection with the blastopore region, either through closure of the blastopore (which often occurs in inhibition) or by actual rupture and so is carried apically and remains closed posteriorly, and the rectal region is absent. In short these larval


LARVAL DEVELOPMENT IN THE SEA URCHIN 81

forms show differential acclimation in the more apical regions and differential inhibition in the more basal.

Figures 40 to 46 show the final stages of differential acclimation in alcohol after still greater degrees of inhibition. In all figures the apical end is uppermost, but the antero-posterior axis has been obliterated to such an extent that in most cases it is impossible to distinguish anterior and posterior regions. The most apical portion of the ciliated band is differentiated in figures 40 to 43, but in figures 44 to 46 differentiation ceases at an earlier stage. The different positions, degrees of differentiation, and relations of the entoderm depend upon the relations between the time when apical acclimation began, upon the degree of differentiation of the entoderm and upon the degree of direct inhibition. Where the direct inhibition does not bring about closure of the blastopore (fig. 15), the entoderm may separate between stomach-intestine and rectal regions, the latter remaining at the basal end and undergoing degeneration (fig. 40), or elongating and maintaining its normal basal connection (fig. 41). In other cases the blastopore closes completely, and the entoderm loses connection with the blastopore region, but remains in contact with the apical region of the body wall, and is carried with it when elongation or enlargement occur. Under these conditions it may differentiate completely, or the rectal region (fig. 40), or mouth and oesophagus (figs. 41, 42) may be absent, or lastly, it may remain a rounded vesicle (figs. 43 to 46).

Where the entoderm remains a closed vesicle it often undergoes degeneration after a few days (figs. 47, 48) as in differential inhibition (p. 74), leaving anenteric forms which may remain alive for several days longer.

Even though no skeleton develops in these forms, more or less aggregation of mesenchyme in the basal region occurs as in normal development, while, in the more extreme degrees of inhibition without acclimation, the mesenchyme remains irregularly scattered in the blastocoel. Evidently in acclimation the apico-basal metabolic differences are sufficient to constitute effective factors in determining the localization of the mesenchyme, while a sufficient degree of inhibition makes them ineffective.

JOURNAL OF MORPHOLOGY, VOL. 28, NO. 1


82 C. M. CHILD

If this localization is chemotactically determined, as is commonly believed, then we may say that a certain difference in metabolic rate along the apico-basal axis is necessary as a basis for the differential chemotactic action.

Attention must be called to the possibility that, in some of the forms shown in figures 40 to 48, the original apical region has been killed before acclimation, and that the apical outgrowth is a reconstitution from more basal regions. It is perfectly certain from the observations on earlier stages that forms like figures 40 to 43 may develop without any apical losses, but it is probable that at least in some cases forms like figures 44, 46 have lost some apical cells. In forms such as figures 38, 39, 40 to 48, which are very characteristic of acclimation in alcohol, the acclimation involves chiefly the more apical regions of the body, while in the basal regions the effect of inhibition persists. The difference between these forms and the acid forms (fig . 41 to 37) is due to the fact that in acids differential acclimation begins earlier in the regions of highest metabolic rate of the various axes and is more complete than in alcohol, therefore the extremes of the resulting form changes are greater than in alcohol. In alcohol differential acclimation overcompensates the effects of differential inhibition only in the more apical regions of the apico-basal axis, and differential acclimation along the antero-posterior axis plays little or no part in determining the form, while in acids the differential effect is merely greater in the apico-basal axis and is also marked in the antero-posterior and medio-lateral directions.

The lower limits of differential acclimation are reached in the partial basal forms where a considerable portion of the apical region of the ectoderm has been killed by the direct inhibiting action of the agent. Without acclimation, such forms remain spherical, the blastopore usually closes and the entoderm becomes a closed vesicle in the blastocoel (figs. 19 to 21), but, under conditions which permit some degree of acclimation, they give rise to forms like figures 49 to 53. In these a small apical outgrowth may occur with the mouth at its base, presumably on its anterior side (fig. 49) ; the mouth may lie at the apex of


LARVAL DEVELOPMENT IN THE SEA URCHIN 83

an apical elongated region (fig. 50); an apical outgrowth may occur and mouth and oesophagus fail to differentiate (fig. 51); or mouth and oesophagus may develop in the absence of an apical outgrowth (figs. 52, 53). Usually in these cases theenteron is not connected with the blastopore region, and the rectal portion does not differentiate (figs. 49 to 51, 53), but occasionally in acids complete differentiation of the entoderm has been observed (fig. 52). Except for the case of figure 52, which has been seen only in acclimation to acids, these forms occur in both alcohol and acids.

In these cases of limited acclimation following a rather high degree of differential inhibition (figs. 40'to 53), the antero-posterior axis and bilaterality have been almost or quite obliterated as effective factors in development and, except for the position of the mouth in some cases (figs. 40 and 49), do not appear in differential acclimation. The larvae remain almost completely radially symmetrical and move with the apical end in advance as long as they live. Only the apico-basal axis remains effective, and the metabolic gradient, which constitutes this axis, has been levelled down to such an extent that it is much less effective in determining and localizing differentiation than under the usual conditions. These partial acclimations, as well as the more extreme differential inhibitions, show very clearly that the minor axial gradients can be leveled down and obliterated to such an extent that they do not reappear, while the apico-basal gradient may still remain effective to some extent. This can only mean that these minor metabolic gradients are less permanently recorded in the protoplasmic substratum (Child, '15 c, pp. 33-35), and this in turn must mean that the high ends of these minor gradients represent lower metabolic levels than the high end of the apico-basal gradient, i.e., the apical region. This conclusion is in agreement with all the facts. If the apical region is the region of highest metabolic rate in the individual, it follows of course that the high ends of gradients in other directions must have a lower rate than the high end of the apicobasal gradient; moreover, the death gradient is much more distinct in the apico-basal axis than in other directions (Child,


84 C. M. CHILD

'16 c), and the minor axes are less effective as factors in localization and differentiation, or become effective later than the major axis. This differential obliteration of axial gradients is one of the most significant results of these experiments, and the forms in which only the apico-basal gradient remains as an effective factor in development approach in certain respects the more primitive types of echinoderm larvae.

THE FORMS RESULTING FROM DIFFERENTIAL RECOVERY

In recovery after temporary action of an inhibiting agent, a distinction must be made between the general and the differential effect. Where the effect of the inhibiting agent has not been so great that recovery is impossible, all parts of the body undergo an increase in metabolic rate after return to water. This is the general effect. The differential effect appears in the differences in rate and degree of recovery along the axes and is like the differential effect in acclimation; i.e., the higher the metabolic rate, the more rapid and complete the recovery, but since in the developmental stages of the sea urchin recovery, like acclimation, follows and is superimposed on the differential effects of inhibition, the forms resulting from differential recovery, like those resulting from differential acclimation, may show various combinations of differential inhibition and differential recovery. Since recovery involves not merely the differential changes in metabolic rate, but a great increase in rate in all parts after removal of the inhibiting agent, development may proceed farther, if the temporary inhibiting action be not too great, than where the agent is present throughout. This difference is responsible for certain interesting features of recovery which will be considered below.

Unfortunately the data concerning recovery are not complete. The close of the breeding season of Arbacia put an end to the experiments when only KCN and alcohol had been used in this way, but, since the results with these agents are similar to the results of differential acclimation, there is every reason to believe that the same would be true for alkalies and acids.


LARVAL DEVELOPMENT IN THE SEA URCHIN 85

Where inhibition is slight and limited to the earlier stages of development, differential recovery may appear as elongation of the apical region in the gastrula (figs. 54, 55) or in prepluteus stages (fig. 56). Such forms give rise to plutei like the acclimation plutei of figures 23 and 24 with very large and long oral lobes, wide angles of divergence, and short body.

From forms of this sort, the gradation, through various combinations of differential inhibition and differential recovery, to forms which show no differential recovery is complete. Certain features of these forms require some consideration.

Where the inhibition is so great that minor gradients are almost or quite obliterated, while the apico-basal gradient still remains an effective factor in development, recovery produces a series of forms resembling the differential acclimation forms of figures 37 and 40 to 53. Figures 57 to 73 show examples of differential recovery after a considerable degree of inhibition by cyanide. In all cases the minor axes are almost or quite obliterated, but there is usually elongation in the apico-basal axis, and in many cases (figs. 57 to 64) an apical outgrowth, a rudimentary oral lobe, on which the apical portion o