Book - Experimental Embryology (1909)
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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 II Cell-Division And Growth
1. Ce1l-division 2. Growth
1. Urrnvitation
2. Mechanical agitation
Electricity and magnetism
Light
Heat
Atmospheric pressure. The respiration of the embryo.
Osmotic pressure. The role of water in growth
The chemical composition of the medium
Summary
(1) The initial structure of the germ as a cause of differentiation.
1. The modern form of the prefurmationist doctrine 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
Chapter V Driesch’s Theories Of Development - General Reflections And Conclusions
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
APPENDIX B
On the part played by the nucleus in (lifferenti:L’tion
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.
s of 88,
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.
Cite this page: Hill, M.A. (2024, June 25) Embryology Book - Experimental Embryology (1909). Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_Experimental_Embryology_(1909)
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