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==Contents==
==Contents==
[[Book - Experimental Embryology (1909) 1|Chapter I Introductory]]
[[Book - Experimental Embryology (1909) 1|Chapter I Introductory]]


[[Book - Experimental Embryology (1909) 2|Chapter II Cell-Division And Growth]]
[[Book - Experimental Embryology (1909) 2|Chapter II Cell-Division And Growth]]
 
# Ce1l-division
1. Ce1l-division
# Growth
2. Growth


[[Book - Experimental Embryology (1909) 3|Chapter III External Factors]]
[[Book - Experimental Embryology (1909) 3|Chapter III External Factors]]
 
# Grravitation
1. Urrnvitation
# Mechanical agitation
 
# Electricity and magnetism
2. Mechanical agitation
# Light
 
# Heat
Electricity and magnetism
# Atmospheric pressure. The respiration of the embryo.
 
# Osmotic pressure. The role of water in growth
Light
# The chemical composition of the medium
 
# Summary
Heat
 
Atmospheric pressure. The respiration of the embryo.
 
Osmotic pressure. The role of water in growth
 
The chemical composition of the medium
 
Summary


[[Book - Experimental Embryology (1909) 4|Chapter IV Internal Factors]]
[[Book - Experimental Embryology (1909) 4|Chapter IV Internal Factors]]


(1) The initial structure of the germ as a cause of differentiation.
(1) The initial structure of the germ as a cause of differentiation.
# The modern form of the preformationist doctrine
# Amphibia
# Pisces
# Amphioxus
# Coe-lenterata
# Ecliinodcrmata
# Nemertinen
# Ctenophora
# Chaetopoda and Mollusca
# Ascidia
# General considerations and conclusions
# The part played by the spermatozoon in the determination of egg-strucure
# The part played by the nucleus in differentiation


1. The modern form of the prefurmationist doctrine
(2) The actions of the parts of the developing organism on one another
2. Amphibia
3. Pisces
4. Amphioxus
5. Coe-lenterata
6. Ecliinodcrmata
7. Nemertinen . . . . . . . . 204
8. (.‘tenopho1':i . . . . . . . . 208
9. Chaetopoda and Mollusca . . . . . . 213
10. Ascidia . . . . . . . . . 229
11. General consiileratious and conclusions . . . 240
12. The part. played by the spernmtozoon in the determination of egg-.<ztructm'e . . . . . 247
13. The part played by the nucleus in ilifl'e1'enti;iti0n . . 251
(2) The actions of the parts of the developing oiganism on
one another 271


[[Book - Experimental Embryology (1909) 5|Chapter V Driesch’s Theories Of Development - General Reflections And Conclusions]]
[[Book - Experimental Embryology (1909) 5|Chapter V Driesch’s Theories Of Development - General Reflections And Conclusions]]
Line 109: Line 97:


APPENDIX A
APPENDIX A
On the .’~y)1)l)lL'l2l'y of the egg, the symmetly of scglnentation, and the symmetry of the embryo in the Frog  
On the symmetry of the egg, the symmetry of segmentation, and the symmetry of the embryo in the Frog  
 


APPENDIX B
APPENDIX B


On the part played by the nucleus in (lifferenti:L’tion
On the part played by the nucleus in differentiation
 
I.\'m<;x or AUTl{0I{
 
Ixmzx or SUBJPJCTS
 
ADDENDA
 
==Appendix B==
 
ON THE PART PLAYED BY THE NUCLEUS IN DIFFERENTIATION
 
(i) BOVERI has more recently (Zellen-Studim, vi, Jena, 1907)
published a very elaborate account of the irregularities produced
by dispermy in Echinoid eggs, in which are brought forward
 
still more facts in proof of the qualitative difference of the
chromosomes.
 
As has been stated above, p. 263, dispermy is induced by
the simple expedient of adding a large quantity of sperm to the
eggs. The following types of dispermy are distinguished.
 
A. Tetracentric, i. e. each sperm centre divides.
(i) 'I‘etraster, with four spindles.
 
(ii) Double spindle, i. e. the female and one male pronucleus
lie in one spindle, the other male lies aside in its spindle.
 
B. Tricentric, one sperm centre remaining undivided.
(i) Triaster, a tripolar figure with three spindles.
 
(ii) Monaster-amphiaster, the undivided sperm centre remaining apart with one sperm nucleus.
 
C. Dicentric, neither sperm centre dividing.
(i) Amphiaster, a spindle is formed between the two centres.
 
(ii) Double monaster: the centres remain apart, one with
one male, the other with the other male and the female
pronucleus.
 
The segmentation of these eggs is as follows.
 
The tetraster divides simultaneously into four, which may
either lie in one plane if the divisions are meridional, or be tetrahedrally arranged. In the first case another meridional division
ensues, followed by an equatorial, then ‘eight micromeres are
formed, eight macromeres, and sixteen mesomeres. In the latter
case not more than three cells can share in the micromere region
and only four or six of these are produced. The triaster eggs,
having divided simultaneously into three (meridionally), subsequently show six micromeres, six macromeres, and twelve
mesomeres.
 
The segmentation of the double spindle eggs is interesting and
important. Usually the egg divides across the two spindles
312 APPENDIX B
 
into two binucleate cells, but it may divide at once into four, or
into three, one of which is binucleate. The interest lies in the
binucleate cells, for they continue to produce uni-nucleate and binucleate cells until the latter divide simultaneously into four,
and this simultaneous division may sometimes involve an irregular
distribution of the chromosomes, with fatal consequences to the
cell. Bovcri had already produced evidence of the evil effects of
an irregular distribution of the 3 n x 2 chromosomes present in
triasters and tetrasters. A more detailed account is now given.
 
Of the tripartite (triaster) ova about 8 % on an average produced Plutei. In these larvae three regions may be distinguished
in the egg by the size of the nuclei (proportional to the number
of chromosomes) and the boundaries between them may be shown
to correspond to the divisions between the three blastomeres.
The form is asymmetrical in skeleton and pigment, but Bovcri
shows that both sides are normal, as though the larva had been
compounded of two types such as occur, as individual variations,
in any culture. It is suggested therefore that the slight differences in the two sides are due to difierences in the two sperms.
 
Some of the larvae have partial defects in skeleton or pigment,
or the skeleton may be much reduced on one side, or one-third of
the cells may be pathological, i. e. disintegrate in the segmentation
cavity, while the remaining two-thirds are sound and sometimes
symmetrical. In this case it is supposed that the degenerate
cells had separated from the others at an early stage, and that
the remainder had had time to recuperate. In others two-thirds
are degenerate, one-third normal, or all three degenerate. When
the three blastomeres are isolated and allowed to develop independently, segmentation is partial, with two micromeres, two
macromeres, and four mesomeres, and often all three develop
normally up to the blastula stage. After that only one or two,
rarely all three, become Plutei, the rest giving rise to stereoblastulae or stereogastrulae, full of degenerating cells.
 
The isolated quarters of tetrasters also segment partially
and normally, but few give rise to Plutei. The whole simultaneously quadripartite eggs only rarely give rise to what may be
called a Pluteus (2 cases in 1500) ; but very degenerate larvae
are found, with masses of disintegrating cells inside, which are
assigned to one of the four blastomeres. Stereogastrulae-—with
nuclei of all the same size--are frequent.
 
As has been alread mentioned, Bovcri points out that the
probability of each cell’ of a triaster receiving a complete set of
the 71. chromosomes of the species when there are 3 n x_2 to be distributed must be greater than‘ that of each cell of tetraster
obtaining a full complement, and the probability for one isolated cell must be greater than that for the whole egg. What the
mathematical values of these probabilities are Boveri does not
know, though he makes an attempt to reckon them—not
theoretically, but by means of a mechanical apparatus; the
attempt is not quite successful. The fact, however, remains that
eight per cent. of the triasters produce normal Plutei, only -06 per
cent. of the tetrasters. This does not depend on the cells receiving
too much or too little chromatin (see p. 265), nor again on the
fact that the ratio between size of nucleus and size of cytoplasm
(see pp. 268, 269) can only be satisfied by certain definite
numbers of chromosomes, and the only explanation remaining is
that for normal development of each and every part the nucleus of
each cell must contain a complete set of the specific chrosomomes ;
from which it follows that the chromosomes are qualitatively
unlike.
 
A word may be said about the double-spindled eggs (Type
A. i). The larvae from these sometimes show abnormal regions,
and this is attributed to one or more of the binucleate cells
having divided with a tetraster and irregular distribution of
chromosomes. Of all such eggs 50 % gave rise to normal Plutei.
 
The degenerative changes undergone by the nuclei of these
larvae are of several types, to be associated again with differences
in the combinations of chromosomes.
 
(ii) Boveri’s experimental proof of the qualitative difference
of the chromosomes does not of course of itself involve a belief
in the individuality of these bodies, for if the chromatin is
concerned in inheritance, it is necessary to suppose that the
number of qualitatively distinct bodies is far greater than the
number of chromosomes, and these bodies may be differently
grouped during each successive resting stage.
 
The hypothesis of the individuality of the chromosomes, i.e. of
a constancy in the manner of grouping of these particles, rests
in the first instance on such facts as those observed by Sutton in
B2-ac/:3/stola, where in the spermatogonia the chromosomes are of
dilferent sizes, which may however be arranged in pairs, together
with an odd one or accessory chromosome. 1 In the resting stage
the accessory chromosome remains apart in a separate vesicle,
while the large chromosomes lie in separate pockets of the
nuclear membrane, the small ones, each as a separate reticulum,
in the main body of the nucleus. In the spermatocyte a number
of bivalent spiremes appear, which show the same dilferences of
sizes a the pairs of chromosomes previously, and the accessory
chromosome.
 
The accessory chromosome passes into two only of the four
spermatids and is supposed to be a sex-determinant.
 
 
Similar facts have been reported by Wilson for several Insects
(see Joum. Esp. Zool. ii, iii, 1905, 1906). '
 
Wilson finds constant size differences between pairs of chromosomes, and either an accessory odd chromosome (which passes
into only one half of the germ cells) or a pair of idio-chromosomes of unequal size (one of which goes to one half, the other to
the other half of the spermatozoa), or both the accessory and the
idio-chromosomes (giving four kinds of spermatozoa). The idiochromosomes are supposed, again, to play a part in sex-determination. Several other observers have found these accessory
chromosomes, idio-chromosomes, and pairs of chromosomes of
difierent sizes in various Insects (Boring, Journ. E211. Zool. iv.
1907 ; Stevens, ibid. ii. 1905, v. 1908; McClung, Biol. Bull. iii.
1902, ix. 1905; Montgomery, Biol. Bull., vi. 1904; Baumgartner, Biol. Bull. viii. 1904-5 ,- Zweiger, Zool. Anz. xxx. 1906;
Nowlin, Jomw. Exp. Zool. iii. 1906); in Spiders (Wallace, Biol.
Bull. viii. 1904»-5 ; Berry, Biol. Bull. xi. 1906); and in Myriapods (Blackman, Biol. Bull. v. 1903 ; Medes, Biol. Bull.
ix. 1905).
 
It is a noteworthy fact that the accessory chromosome retains
its individuality in the resting stage (looking like a chromatin
nucleolus), while the others break up. The belief in the individuality of these others rests therefore on the constancy of the relative sizes from generation to generation.
 
Further support for the hypothesis may be derived from theoretical speculations. VVe know that only 2; (one-half the normal
number) chromosomes are necessary for normal development
provided that they comprise a complete set. In sexual reproduction n maternal unite with n paternal. A study of the reducing division shows that 1: whole chromosomes first pair with
and are then separated from or whole chromosomes, and that
when they dilfer in size those of the same size pair together, and
it looks as though paternal were here separated from maternal,
though the distribution of paternal and maternal to the two cells
will difier, almost certainly, in diiferent cases.
 
If the particles of which the chromosomes are composed are
also to be paired and separated, it would appear to be necessary
that their groupin should be constant, in other words that the
chromosomes shou d retain their individuality.
 
(iii) A case of heterogeneous fertilization between eggs of Seaurchins and the sperm of Anletlon has been described above
(p. 262). Loeb has recently succeeded in rearing Plutei from
the eggs of Slrongylocmlrolue fertilized by the sperm of a
Mollusc (0/lloroaloma). Cytological details are not given (Arc/E.
Eul. Mecfi. xxvi. 1908). ‘
 
 
==Index Of Authors==
 
Agassiz: effects of fertilization in Ctenophors, 250.
 
Aristotle: theory of development, 13.
 
— the soul in function and development, 292 sqq.
 
— mechanism and teleology, 296.
 
Auerbach :' segmentation of Ascuris
nigrovenosa, 33.
 
von Baer, 16.
 
Balfour: effect of yolk on segmentation, 29, 88.
 
Bataillon: monstrosities
osmotic pressure, 120, 135.
 
—- artificial parthenogenesis, 124.
 
Bergh: cell-division in germ-bands
of Crustacea, 34.
 
Berthold: surface-tension and celldivision, 41, 42.
 
Bischofl‘, 16.
 
Blane: effect of light upon the
development of the Chick, 94, 96.
 
Boas: rate of growth in man, 63.
 
— change of variability, 73, 74.
 
— diminution of correlation coefiicient, 75.
 
Bonnet : emboitement, 14.
 
— preformation, 15.
Bonnevie : diminution of chromosomes in Ascaris lumbricoidcs, 258.
Born : gravity and development, 18,
88-85.
 
— pressure experiments on Frogs’
eggs, 34, 35.
 
Boveri : early development of Slrongylocentrotus, 23, 183-185.
 
— egg of Strongylocentrotus stretched,
39.
 
— suppression of micromeres in
Strongylocentrotus, 186.
 
-— causes of the pattern of segmentation, 197.
 
— karyokinetic plane, sperm path,
:11 ng first furrow in Strongylocentrotus,
 
8 .
 
— potentialities of? animal and vegetative cells, 192.
 
— stratification of cytoplasmic substances, 242, 280.
 
-- characters dependent on cytonlmam in Flnhinnid larvae. 261.
 
due to
 
Boveri : diminution of chromosomes
in Ascaris megalocephala, 252, 255-257.
 
— due to a difference in the cytoplasm, 257.
 
— hybrid larva from enucleate egg
fragment with characters of male
parent, 253, 258-260.
 
— irregular distribution of chromosomes a cause of abnormality, 253,
263-266.
 
— individuality of chromosomes and
chromatin, 256, 263.
 
—part played by nucleus in differentiation, 266, 285.
 
—possiblo significance of reducing
divisions, 266.
 
— number of chromosomes, size of
nucleus, and size of cell, 68, 267,
268.
 
—2méclear division not qualitative,
 
6 .
 
Bowditch: rate of growth in man,
63.
 
-- change of variability, 73.
 
Brauer : Branchipus, 22, 24.
 
Brooks: Lucifer, 22.
 
de Butfon : Preformation, 15.
 
Bullzt: artificial parthenogenesis,
12 .
 
Bumpus: change of variability in
Litlorina, 71, 72.
 
Bunge: respiration of Ascaris, 112.
 
Castle : see Davenpofl: and Castle.
 
Chabry: segmentation furrows and
embryonic axes in Ascidians, 229.
 
—- development of isolated blastemeres in Ascidians, 229, 230.
 
Child : critique of Driesch’s vitalism,
292, note.
 
Chun : isolated blastomeres of Ctenophora, 209.
 
Conklin: maturation, fertilization,
and development of Cynthia, 230236.
 
— development of isolated blastemeres in Oyntlzia, 237.
 
— development of pieces of gastrula
in Cynthia, 238.
 
— streaming movements of protonlnsm. 40.
316 INDEX OF
 
Crampton : isolated blastomeres of
Ilycmesaa, 215, 216.
 
— efieot of removal of the polar lobe,
217.
 
Dareste: mechanical agitation of the
Hen’s egg, 89.
 
— electricity, 91.
 
Davenport : catalogue of ontogenetic
processes, 4 sqq.
 
— definition of growth, 58.
 
— rate of growth, 69.
 
— the role of water in growth, 58,
59, 115, 116.
 
- and Castle : acclimatization of eggs
of Bufo to heat, 100.
 
Delage : causes of artificial parthenogenesis, 124.
 
-- number of chromosomes in artificial parthenogenesis and in merogony, 125.
De Vries : importance of potassium
for turgor of plant-cells, 146.
 
Doncaster: hybrid Echinoid larvae,
26].
 
Driesch: effect of light in development, 94.
 
— abnormal segmentation in Erhinus
produced by heat, 105.
 
— Anenteria, produced by heat,
106.
 
—- segmentation made irregular by
dilution of sea-water, 118.
 
—— pressure experiments on Echinoid
eggs, 37, 38, 185, 240.
 
—- cell-division suppressed by pressure and dilute sea-water, 55; and
by heat, 105.
 
—nuclear division not qualitative,
186.
 
— blastomeres disarranged, 187, 188.
 
— isolated blastomeres of Echinoids,
190, 191, 193, 194.
 
— potentialities of animal and vegetative cells, 193, 194, 201, 242, 243.
 
— fragments of blastulae and gastrulae in Echinoderms, 194.
 
— potentialities of ectoderm and
agghenteron, and their limitations,
1 .
 
— development of egg fragments of
Echinoids, 195, 196.
 
— germinal value, surface-area of
larvae, and number of cells, 197199, 269.
 
— one larva from two blastulae, 202.
 
— and Morgan : isolated blastomeres
of Ctenophora, 210, 211.
 
—2e1gg-fragments of Ctenophora, 30,
 
2!
 
AUTHORS
 
Drgggchz development of Myzostoma,
 
— isolated blastomeres and parts of
larvae in Phallusia, 288, 289.
 
— first furrow and sagittal plane in
Echinoids, 250.
 
— characters which depend on cytoplasm in Echinoid larvae, 261, 262.
 
— number of organ-forming substances in cytoplasm, 246, 284,
286.
 
—— theory of egg-structure, 281, 286,
292.
 
— reason for limitation of potentialities, 192-194, 201, 212, 242, 243,
281, 282, 284, 291.
 
--fate a function of position, 188,
282.
 
—- return of displaced mesenchyme
cells in Echinus, 274.
 
- stimuli in ontogeny, 20, 277, 28"284.
 
— part played by nucleus in differentiation, 266, 284, 285.
 
—— equipotential and inequipotentiul
systems, 176, 277, 285.
 
— rhythm of development, 3.
 
—- harmony of development, 284.
 
—- composition in development, 3,
285.
 
— self-difierentiation, 284.
 
—- teleology, static, 286, 291, 292,
297.
 
— —- dynamic, 291, 292, 297.
 
— vitalism, 20, 289 sqq.
 
Edwards : physiological zero for
Home egg, 102.
 
-- growth without differentiation,
104.
 
Endres and Walter : post-generation
of missing half-embryo, 171.
 
Eycleshymer: first furrow
sagittal plane in Necturus, 168.
 
and
 
Fabricius : views on development,
13.
 
Fasola : electric currents, 91.
 
Fehling : growth of the human
embryo, 59, 60, 63.
 
Feré : effect of sound-vibrations upon
the Chick, 90.
 
_ ._ of light, 96.
 
— malformations due to high temperatures, 105. .
 
—- need of oxygen for the Chick, 109.
 
—— monstrosities produced by various
chemical reagents, 18,2.
INDEX OF AUTHORS
 
Fischel, A. : hybrid Echinoid larvae,
261.
 
— variability of Duck embryos, 71.
 
Fischel, H. : isolated blastomeres of
Ctenophora, 210, 211.
 
-— derangement of blastomeres in
Ctenophora, 211.
 
Fischer: artificial parthenogenesis,
124. ’
Foot : polar rings in Allolobophom,
 
251.
 
Garbowski : function of pigment
ring in Strongylocentrotus egg, 192.
— first furrow and sagittal plane in
 
Echinoids, 260.
 
— grafting of blastulae fragments of
Echinus, 202.
 
Gerassimow: size of nucleus and
cells in Spirogyra, 269.
 
Giacomini: need of oxygen for the
Chick, efiect of low atmospheric
pressure, 109, 110.
 
Giardina : difierentiation of chromatin in female cells of Dytiscus.
 
Godlewski : the respiration of the
Frog’s eg, 110, 112, 113.
 
-— heterogeneous cross-fertilization,
262.
 
Graf : fusion of blastomeres, 56.
 
Greeley: artificial parthenogenesis
produced by cold, 108.
 
— low temperatures and absorption
of water, 108.
 
Grobben : Cetochilus, 22.
 
Groom : effect of fertilization in
Cirripedes, 250.
 
Gigiber: regeneration in Protozoa,
 
54.
 
Gurwitsch : monstrosities produced
in Amphibian embryos by chemical
reagents, 120, 123.
 
Hacker : Cyclops, 22.
 
Haeckel: recapitulation, 16.
 
— development of fragments of
blastulao of Crystallodes, 181, note.
Hr;ller : preformation and epigenesis,
 
5.
 
Harvey: epigenesis, 13.
 
— metamorphosis, 14.
 
Hecker: growth of the human embryo, 62, 63.
 
Hansen: growth of guinea-pig embryos, 62.
 
Herbst : potassium, sodium, and
lithium larvae of Echinoderms,
136-140.
 
—- significance of monsters for origin
of variatiops, 141.
 
317
 
Herbst : necessity of elements present
in sea-water for normal development of Echinoid larvae, 141 sqq.
 
—— separation of blastomeres of Seaurchins in calcium-free sea-water,
 
45.
 
— stimuli in ontogeny, 20, 272, 273,
285.
 
— formation of Arthropod blastederm oxygenotactic, 114.
 
—— arms of Plutous due to presence of
skeleton, 187, 138, 144, 149, 274, 275.
 
I-Ierl itzka, development of half-blastomeres of Newt, 173.
 
Hertwig, 0. : centrifugalized Frog’s
egg, 29, 87.
 
—- rules for nuclear and cell division,
31, 32, 85.
 
— — confirmed by pressure experiments, 34-36.
 
— gravity and Echinoderm eggs, 78.
 
—— insemination of Frog's egg, 79.
 
— cardinal temperatures for Rana
 
fusca. and csculenta, 97.
 
— monstrosities produced by high
and by low temperatures, 99.
 
— temperature and rate of development, 100.
 
—— monstrosities produced in Amphibian embryos by sodium chloride,
119, 135.
 
— first furrow and sagittal plane in
Frog's egg, 165.
 
— compressedeggs: disproof of qualitative nuclear division, 34—86, 168,
169, 240.
 
— development of half-blastomere of
Frog’s egg, 169.
 
— mutual interactions of developing parts, 271, 285.
 
Hertwig, 0. and R. : fertilization
processes altered by heat and cold,
107.
 
— — by alkaloids, 126 sqq., 263.
 
His: mechanical explanation of
development, 3.
 
—- germinal localization, 17, 158.
 
— the blastoderm oxygenoti-opic,114.
 
Hunter: artificial parthenogenesis
by concentrated sea-water, 124.
 
Iijima: spiral asters in Nephelis egg,
40.
 
Jenkinson: pressure experiments on
eggs of Antedon, 37, note.
 
— abnormalities of Frog embryos
produced by various solutions not
due to increased osmotic pressure,
120, 133-136.
318
 
Jenkinson: plane of symmetry, first
furrow and sagittal plane in Frog's
egg, 165-168.
 
Jennings: fertilization spindle in
Asplanclma, 34.
 
Kaestner: cardinal temperature
points for the Hen‘s egg, 102.
 
— malformations due to low tem~
peratures, 104. '
 
Kant : teleology, 286-289, 292, 297.
 
Kastschenko: injuries to blastoporic
lip in Elasmobranchs, 178.
 
Kathariner: gravity and the gray
crescent of the Frog's egg, 86.
 
King : cause of differentiation of lens,
276, 276.
 
Knowlton : sec Lillie and Knowlton.
 
Kolliker: 16.
 
Kopsch : first furrow and sagittal
plane in Frog's egg, 165, 168.
 
—— efl'ect of injuries to blastoporic lip,
178.
 
Korschelt: fusion of ova in Ophryotmcha, 202.
 
— nucleus of egg-cell in Dyfiscus, 252. .
 
Kostanecki and Wierzejski: efi'ect of
fertilization in Physa, 250.
 
Kowalewsky: 16.
 
Kraus : the role of water
growth of plants, 58.
 
Lang : effect of fertilization in Polyclads, 250.
 
Leibnitz : preformation, 15.
 
Lewis: causes of formation of lens
and cornea, 275, 276.
Lillie and Knowlton: eflect of low
temperatures in Amphibia, 100.
— temperature and rate of development, 101.
 
Lillie: effects of salts on ciliary
movement, 135.
 
— ghysiologically balanced solutions,
1 6.
 
in the
 
— toxicity and valency, 136.
 
Loeb : suppression of cell-division
in Echinoids and Fishes, 56, 117.
-— eflect of light in development, 94.
—the respiration of Otmolabrua and
 
Fundulua eggs, 111.
 
—— the respiration of the ova of
Echinoids, 112.
 
— function of oxygen in regeneration
of Tubular-ia head and other processes, 114, 278, 274.
 
-— efi'ect of hypertonic solutions on
Fundulus and Arbacia eggs, 117.
 
--exovates produced by dilute seawater, 118, 190, 194, 195.
 
INDEX or AUTHORS
 
Loeb: artificial parthenogenesis,
121, 124.
 
—- etfect of potassium cyanide in prolonging life of ova, 131, 132.
 
— eflect of certain salts on Fundulus
embryos and on Plutei, 135.
 
— toxicity and antitoxicity functions
of valency, 186.
 
-— effect of alkalies, 151.
 
— effect of gravity on Anmmularia,
272, 273.
 
-gégterogeneous cross-fertilization,
 
Lombardini : electric currents, 91.
 
Lyon : need of oxygen for the eggs of
Arbacia, 112.
 
— action of potassium cyanide, 132.
 
Malebranche : preformation, 15.
 
Malpighi: preformation, 14, 15.
 
Marcacci : mechanical agitation of
Hen's eggs, 90.
 
Mark: spiral asters in eggof Lz‘maac,40.
 
Mathews: artificial parthenogenesis
by mechanical agitation, 90.
 
—— effects of atropine and pilocarpine
on Echinoderm eggs, 131.
 
—toxicity and decomposition tension,
136.
 
— see also Wilson (E.B.)and Mathews.
 
Mencl : formation of lensin SaImo,276.
 
Metsclinikoif : separation of blastemeres of Oceania, 181.
 
-—fusion of blastulae in Mitrocoma, 202.
 
Minot : rate of growth defined, 60.
 
—— change of rate of growth of guineapigs, 61.
 
— - of rabbits, 62, 68.
 
— — ofchickens, 67.
 
— coeflicients of growth, 65.
 
— senescence, 65.
 
-- increase of cytoplasm, decrease of
mitotic index, 65.
 
— change of variability in guineapigs, 71. _
— genetic restriction, 246, 277.
Mitrophanow: malformations due to
low and high temperatures, 104.
— necessity of oxygen for the Chick,
109.
 
Moore : sodium sulphate an antidote
to sodium chloride, 135, 186.
 
Morgan : suppression of cell-division
in Arbacia, 56, 118.
 
- gravity and the gray crescent of
the Frog's egg, 86.
 
-— monstrosities produced by low
temperatures in Ranapaluslris, 100.
 
— need of oxygen for the Frog's egg,
110.
INDEX OF AUTHORS
 
Morgan :lithium salts used to produce
alzlgéiormalities in Frog's eggs, 120,
 
— attempts to induce
parthenogenesis, 124.
 
— number of chromosomes in artificial parthenogenesis, 125.
 
— artificial parthenogenesis produced
by cold, 108.
— first furrow, plane of symmetry,
and sagittal plane in Frog's egg,
165,168.
 
— development of half-blastomere of
 
Frpg's egg ; post-generation, 170,
 
17 .
 
— development of vegetative cells of
Frog’s egg, 173.
 
— potentialities of half-blastomeres
in Teleostei, relation of flrstfurrow
tn sagittal plane, effect of removal
of yolk, 178.
 
— effect of injuries to blastoporic lip,
179.
 
— number of cells in partial larvae
of Amphioxus, 181.
 
— potentialities of ectoderm in
Echinoids, 195.
 
— development of egg-fragments of
Echinoids, 197.
 
— number of cells in partial larvae
of Echinoids, 198.
 
— fusion of blastulae of Sphaerechinua,
201.
 
— and Driesch: isolated blastomeres
and egg-fragments of Ctenophora,
210-212.
 
— micromercs of Ctenophore egg, 30.
 
—- characters of hybrid Echinoid
larvae, 260.
 
Moscowski : gravity and the gray
crescent of the Frog's egg, 86.
 
Miihlmann : prenatal growth-rate
in man, 64.
 
artificial
 
Nfigeli : permutations of original
elements in development, 286.
 
Pander: 16.
 
Pearson : variability in man, 73.
 
Pfliiger: isotropy of the cytoplasm,
18, 158.
 
—--influence oi’ gravity on development, 18, 78, 81-83, 168.
 
-- rule for direction of nuclear
division, 32, 85.
 
Plateau : principle of least surfaces,
41, 43.
 
Platnerz 280.
 
Pott : growth of the Chick, 59, 60, 67.
 
319
 
Pott and Preyer: respiration of the
Chick, 112.
— loss of weight of Hen’s egg due to
evaporation from albumen, 115.
Preyer : rate of growth, 60.
 
Quetelet: change of rate of
in man (weight), 68.
 
— — (stature), 69.
 
— — (other dimensions), 90.
 
growth
 
Rauber : efiect of reduced atmospheric pressure on the Frog’s egg,
110.
 
— elfect of pure oxygen on the eggs
and tadpoles of the Frog, 118, 114.
 
Reichert: 16.
 
Remak : 16.
 
Robert : mechanics of spiral segmentation, 45-47.
 
— rate of growth in man, 68.
 
—-— change of variability, 73.
 
Rossi : efi‘ect of electricity on
Amphibian eggs, 91.
 
Roux : aims of experimental embryology, 13.
 
— ‘Mosaik-Theorie ’ of self-differentiation, 17, 158, 279, 286, 297.
 
— qualitative nuclear division abandoned, 19, 159, 240.
 
— idioplasm and reserve-idioplasm,
159, 266.
 
— a half-embryo from one of first
two blastomeres and post-generation of missing half, 159, 162.
 
— coincidence of first furrow and
sagittal plane in Frog's egg, 17, 159,
165. '
 
— the spermatozoon and symmetry
of the Frog's egg and embryo, 80,
165, 247, 248.
 
— meaning of karyokinesis, 252.
 
— dependent diflerentiation, 17, 158,
277, 286.
 
— functional adaptation, 290.
 
-— specific gravity of contents of
Frog’s eg, 79.
 
—- gray crescent of Frog's egg, 80, 165.
 
— influence of gravity on the Frog's
egg, 85-87.
 
— effect of electricity upon the Frog’s
egg, &c., 92.
 
— light and development, 93.
 
— segmentation of Rana esculenta, 26.
 
—- Frog's eggs compressed in small
tubes, 39, 40.
 
— comparison of systems of oil drops
and segmenting ova, 49-58.
 
— cytotropism, 55, 278.
320
 
Roux: cytotaxis, 55.
 
— cytochorismus, 45.
 
-— cytarme, 45, 53.
 
— cytolisthesis, 58.
 
— ‘ Framboisia’, 135.
 
Ruseoni : electric currents, 91.
 
Sachs : law of direction of cell
division, 28.
 
Sala: fertilization processes altered
by cold, 108.
 
- fusion of the eggs of Ascaris, 202.
 
Samassa: effect of pure oxygen at
pressures on the Frog's egg,
 
— effect of lack of oxygen on the
Frog's egg, 119.
 
— effect of various gases on the eggs
of Ascaris, 112.
 
—development of animal cells of
Frog's egg, 173.
 
— Schaper: development of tadpoles
after removal of brain and eyes,
175.
 
—- cause of differentiation of lens,
275.
 
Schulze, F. E. :
Sponges, 22.
Schulze, 0.: gray crescent of Frog’s
 
eg, 80, 247.
 
—— gravity and the Frog’s egg, 86.
 
—- effect of low temperatures on the
Frog's egg, 100.
 
—— first furrow and sagittal plane in
Frog's egg, 165.
 
— double monsters from Frog’s egg,
171.
 
Seeliger : hybrid Echinoderm larvae,
260, 269.
 
Selenka: first furrow and sagittal
plane in Echinoids, 250.
 
Semper: rate of growth in Limnaea, 67.
 
Smith: Peltogaster, 24.
 
Sollmann : after effects of hypertonic
solutions, 124.
 
Spemann : development ofconstricted
Newt's eggs, and embryos, 174, 175.
 
— causes of formation of lens and
cornea, 275, 276.
 
Sumner: injuries to blastoporic lip
of Teleostei, 178, 246.
 
Sutton {individuality of chromosomes
in Brachyslola, 256.
 
Swammerdam : preformation, 14, 15.
 
segmentation of
 
Vejdovsky : unequal centrosomes in
dividing pole-cells, 31.
 
— polar rings in Rhym.-hclmis, 251.
 
Vernon: rate of growth in Strongmlocmtrotus, 67, 70.
 
INDEX or AUTHORS
 
Vernon : alteration of variability in
Echinoid larvae, 71, 74.
 
-— effect of light on Echinoid larvae,
95, 96. '
 
— effects of change of temperature
on Echinoid larvae, 106, 107.
 
-— change of variability produced
by heat, 107.
 
— and by chemical agency, 141, 156.
 
—poisonousness of carbon dioxide
to Sea-urchin eggs, 112.
 
— characters of hybrid Echinoid
larvae, 261.
 
Verworn : behaviour of Protozoa in
an electric current, 93.
 
— regeneration in Protozoa, 254,
note.
 
Walter, sec Endres and Walter.
 
Weber : law of stimuli, 272.
 
Weismann: qualitative
division, 19, 297.
 
— idioplasm, and reserve—idioplasm,
159.
 
Weldon : growth-rate in Carcinus, 71.
 
— change of variability in Carcinus,
72.
 
— — in Clausilia, 73.
 
Wetzel : double monsters
Frog’s egg, 172, 245.
 
Whitman : polar rings in Clepsine,
251.
 
Wierzejski, see
Wierzejski, 250.
 
Wilson, 0. B. : malformations of
Amphibian embryos, 120.
 
— acclimatizution to salt-solution,
136.
 
Wilson, E. B. :
phioxus, 26.
 
—— segmentation of Renilla, 55, note.
 
— unequal centrosomes in dividing
pole-cells, 31.
 
—pressure experiments on eggs of
Nareis, 39, 213, 240.
 
- cytology of artificial parthenogenesis, 124.
 
— development of isolated blastemeres in Amphioxus, 179, 180.
 
—— isolated blastomeres of Oerebratulus,
and fragments of blastulae, 205,
206.
 
— isolated blastomeres of Patella,
218-222.
 
—- of Dentalium, 225, 226.
 
—— removal of polar lobe, 224.
 
— effect of fertilization, 222, 223.
 
— development of egg-fragments,
226, 227.
 
nuclear
 
from
 
Kostanecki and segmentation of Am
 
Wilson (E. B.) and Mathews : spermpath, egg axis, fix-st furrow, and
embryonic axes of Toacopneustes,
185, 249, 250. ‘
 
Windle: effect of magnetism and
electricity on development, 91.
 
Wolff : epigenesis, 16. '
 
Yatgu: egg-fragments of Cerebratulus,
 
2 7.
 
Yung: effect of light on tadpoles,
etc., 94.
 
Zeleny : egg-fragments of Cerebratulus,
206, 207.
 
Zelinka : fertilization
Callidma, 34.
 
spindle in
 
Jnxntsonr’ Y
 
Ziegler : heterodynamic centrosomes, 80.
 
.— formation of micromeres in Cteno
phora, 209, note.
 
-— pressure experiments on egg
gaéiinoids and Ctenophora,
 
— fertilization of Diplogaster, 84.
 
— egg and embryonic axes, 250.
 
Zoja : isolated blastomeres of Hydromedusae, 181, 182.
 
—— animal and vegetative cells of
Strongylocentrotus, 198.
 
Zur Strassen : segmentation of
Asoaiis, 81.


— fusion of the eggs of Ascaris.
Index of Authors


s of
Addenda
88,


==Addenda Et Corrigenda==
==Addenda Et Corrigenda==

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Jenkinson JW. Experimental Embryology. (1909) Claredon Press, Oxford.

Jenkinson (1909): 1 Introductory | 2 Cell-Division and Growth | 3 External Factors | 4 Internal Factors | 5 Driesch’s Theories - General Conclusions | 6 Appendices
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Experimental Embryology

Experimental Embryoijogy

By

J. W. Jenkinson. M.A.. D.Sc.

Lecturer in Embryology in the University of Oxford

(1909)

Preface

For the biologist there are, I conceive, in the main two problems. One is to give an account of those activities or functions by means of which an organism maintains its specific form in an environment. The other is to find the causes which determine the production of that form, whether in the race or in the individual. The solution of the first of these problems is the business of physiology, in the usual sense of the term. The second falls to morphology.


It is with the origin of form that we are here concerned, and in particular with its origin in the individual. The endeavour to discover by experiment the causes of this process — as distinct from the mere description of the process - is a comparatively new branch of biological science, for Experimental Embryology, or, as some prefer to call it, the Mechanics of Development (Entwicklungsmechanik), or the Physiology of Development, really dates from Roux's production of a half-embryo from a. half-blaatomere, and the consequent formulation of the ‘ Mosaik-Theorie’ of self-differentiation. That hypothesis has been the focus of much fruitful criticism and controversy, the experiment has been followed by many others of the same kind, and the present volume is an attempt to sketch the progress of these researches and speculations on the nature and essence of differentiation, as well as of those which deal with growth, cell-division, and the external conditions of development.


In writing this review I have had the very great advantage of an excellent model in the textbook of Korsehelt and Heider (Lehrbuch cler fucrgleichemleat Entwio/cluugsgeschiclzte (Zer 1ve'rbelZo.~e'n T/u'c7'e, Allgemeiner Theil, Jena, 1902). I have indeed followed the general arrangement adopted by these authors fairly closely except in one respect. I believe so strongly that the processes of growth and cell-division, though they always (in the Metazoa) accompany, are yet distinct from, differentiation, that I have felt justified in treating them in a chapter apart from the other internal factors of development. The external factors—whether of growth, celhdivision, or differentiation - are discussed in Chapter III, and the ground is thus cleared for a consideration of the real problem — the differentiation of specific form.

The last chapter is devoted to the theories, scientific and philosophical, of Hans Driesch. I sincerely hope that Herr Driesch will allow my great admiration for the former to atone in some measure for my inability to accept the tenets of nee-vitalism.


It is a very great pleasure to me to acknowledge my indebtedness to the Delegates and Secretaries of the Clarendon Press, and in particular to Professor Osler, for undertaking the publication of this book, as well as for the pains which have been expended in its preparation. Dr. Osler also took the trouble to read through the whole of the manuscript, and Mr. G. V. Smith and Dr. Haldane have been kind enough to look through certain chapters.


To Dr. Ramsden I am under great obligations for his assistance in that part of Chapter II, Section 1, in which surface-tensions are discussed; to Dr. Vernon for calling my attention to Roberts’s work on Anthropometry, and to Mr. Grosvenor for the information embodied in the foot-note on p. 89. Mr. A. D. Lindsay has given me invaluable assistance in those sections of Chapter V which deal with the philosophy of Kant, while, for Aristotle, I was fortunately able to attend Professor Bywater’s lectures on the De Anima.

I can hardly express the debt I owe to Mr. J. A. Smith for much friendly counsel and criticism, although he is, of course, in no way responsible for the philosophical speculations in which I have ventured to indulge.


The illustrations are largely borrowed from Korschelt and Heider’s work, and I must thank Herr Gustav Fischer, of Jena, for his readiness in supplying the blocks. Others are from the original publications‘, and I am obliged to the proprietors for permission to make use of them. A few are my own.


In the appendices will he found an account of some recent work on the relation between the symmetry of the egg and that of the embryo in the Frog, and on the part played by the nucleus in ditt'c1-entiation.

Proceedings of the Boston Society of Natural History, the Journal of Experimental Zoology (Williams 8; Wilkins, Baltimore), the Anm'ir(rn Journal of I‘hysz'ulo_'/_I/ (Ginn & C0., Boston), ZeIIrn~Sfu(Iim (Fischer, Jena), l’erhamIlmI_r/en 410;" A/mlumis-1-hm G(‘.s'¢'”N(‘7I((fl (Fischer, Jena), Er;/cbnisse fiber din Ii'on.m'tzm'ou dcr cIu'onmta'scIzm Kernsubslmz: (Fischer, Jena), .[r¢-kin fiir mik)'osk0])i.s¢*7¢1: .»lm¢tomi(' (Cohen, Bonn), Archizv ff/"r Entwiclcluuysnwvlzanik (Engelinunn, Leipzig), and the Popular Science .llontM3/ (Appleton & Co., New York).

Contents

Chapter I Introductory

Chapter II Cell-Division And Growth

  1. Ce1l-division
  2. Growth

Chapter III External Factors

  1. Grravitation
  2. Mechanical agitation
  3. Electricity and magnetism
  4. Light
  5. Heat
  6. Atmospheric pressure. The respiration of the embryo.
  7. Osmotic pressure. The role of water in growth
  8. The chemical composition of the medium
  9. Summary

Chapter IV Internal Factors

(1) The initial structure of the germ as a cause of differentiation.

  1. The modern form of the preformationist doctrine
  2. Amphibia
  3. Pisces
  4. Amphioxus
  5. Coe-lenterata
  6. Ecliinodcrmata
  7. Nemertinen
  8. Ctenophora
  9. Chaetopoda and Mollusca
  10. Ascidia
  11. General considerations and conclusions
  12. The part played by the spermatozoon in the determination of egg-strucure
  13. The part played by the nucleus in differentiation

(2) The actions of the parts of the developing organism on one another

Chapter V Driesch’s Theories Of Development - General Reflections And Conclusions

Appendices

APPENDIX A On the symmetry of the egg, the symmetry of segmentation, and the symmetry of the embryo in the Frog


APPENDIX B

On the part played by the nucleus in differentiation

Index of Authors

Addenda

Addenda Et Corrigenda

P. 5, 5 lines from bottom, for unicellular read multicellular. P. 28, line 10, after irregular, insert and in Triclads.

P. 57. To Literature acid J. Sacns. Die Anordnung den-Zellen in jiingsten Pflanzentheilen, Arb. Bot. Inst. Wurzburg, ii, 1882. _

P. 114. To Literature add G. BUNGE. Weitere Untersuchungen iiber die Athmung der Wiirmer, Zeitsc-hr. physiol. Chem. xiv, 1890.

P. 140, line 22, for prospective potentialities read prospective significanoes.

P. 225, 2 lines from bottom, for is now placed in road has now moved into.

P. 271. To Literature add W. S. Surrox. On the morphology of the chromosome group in Brachyslola magna, Biol. Bull. iv, 1902.

P. 278. To Literature add J. W. Jnxxmsox. On the effect of certain solutions upon the development of the Frog's egg, Arch. Ent. Mech. xxi, 1906.



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