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Full text of "Studies on dimorphism of spermatozoa"

O .'.'„■ ,( J!. 

Sfilim on Dimorphism of Spermatozoa 





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THE UNIVERSITY 
OF ILLINOIS 
LIBRARY 



Digitized by the Internet 4»'Phive 
in 2013 



http://archive.org/details/studiesondimorphOOsena 



STUDIES ON DIMORPHISM OF SPERMATOZOA 



BY 

CHARLES TIMOTHY SENAY 
B.S. Trinity College, 1914 



THESIS 

Submitted in Partial Fulfillment of the Requirements for the 

Degree of 
MASTER OF ARTS 
IN ZOOLOGY 

IN 

THE GRADUATE SCHOOL 

OF THE 

UNIVERSITY OF ILLINOIS 
1915 



UNIVERSITY OF ILLINOIS 
THE GRADUATE SCHOOL 

„_,..;W8,y 5.1 191 5 

I HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY SUPER- 

\TsioN BY cijyyi3Li:s tlMQlM..3MAY.*. „ 

ENTITLED SIL'2IES....01l-..LIlIGHPiiim...DE....SP.])IB]^.GZDA« 



BE ACCEPTED AS FULFILLING THIS PART OF THE REQUIREMENTS FOR THE 
DEGREE OF IIAST}i:R....OS....ARlS.. „ 




Recommendation concurred in :* 




Committee 
on 

Final Examination* 



^Required for doctor's degree but not for master's. 



UlUC 



STUDIES ON DIMORPHISM OF SPERMATOZOA 
Contents 



I. INTRODUCTION ^^S® 

1. Object of Research 1 

2. Theoretical Basia 1 

3. Listed Species having Sex Chromosomes 2 

4. Previous Work on Size Dimorphism 15 

II. MATERIAL AND METHODS 

1. Most Desirable Species 15 

2. Fixation and Staining 16 

3. Method of Measurement 17 

4. Sources of Error 17 

III. DATA 

Heraiptera 

1. Leptoooria trivittatus 21 

2. Reduviolus ferus 21 

3. Corizua lateralus 22 

4. Euschistus variolarius 22 

5. Cosmopepla carnifex 24 

Coleoptera 

6. Passalus cornutus 24 

7. Berosus striatus 25 

Annelida 

8. Heliodrilus caliginosa 25 

IV. DISCUSSION 26 

Acknowledgments 29 

V. SUMJ/IARY 31 

VI., BIBLIOGRAPHY OF SPERMATOGENESIS 33 



31SS20 



I. INTRODUCTION 
Obj ect of Resear ch 
A limited number of caaes of size dimorphism of spermato- 
zoa has been reported. It is desirable to see if size dimorphism 
is a general phenomenon. Therefore the included piece of research 
was undertaken in the aim of determining if there was size dimor- 
phism of spermatozoa from normal adult individuals of species 
hitherto unstudied. 

3. Theoreti cal Basis 

Numerous investigators have directed their attention to 
the problems of spermatogenesis and oogenesis as they manifest them 
selves in the various groups of animals. As a result of their 
investigations a vast amount of data has been accumulated, the 
large majority of which indicates that there are two kinds of 
spermatids in practically all species, differing quantitatively in 
chromatin content. This chromosomal difference proclaims itself 
in two ways; as the familiar unpaired "x" chromosome and as the 
unequal "x, y" pair of chromosomes. Modifications of these main 
types are found and usually arise by division of the "sex" chromo- 
some, into more numerous units. Examples of all these variations 
will be found in an included list. 

If, as the above evidence indicates, the chromatin con- 
tent of the primitive germ cell is unequally divided in the sperma- 
tids and this division is constant within a given species, then 
there should be spermatozoa of two lengths in the male sexual 
product. Since each of these spermatozoa will be the mean of a 
Gaussian variation curve, a number of other lengths will be found. 



3- 



A graph of the obtained results should be a combination of two 
normal variation curves, with two high points, indicating the two 
means. In cases where there has been no observed difference in the 
spermatids or where the female gametes possess the unequal chrom- 
atin distribution, a simple variation curve might be obtained. 

In the following data "n" indicates the somatic number 
of aatoeomes. "x"^ or "x" and "y" indicate the sex chromosomes. 

3' Lis ted Species haying Sex Chrqmo s ome s 
TYPE I. n/2, n/2 + x. 
Amphibia 

Alytes obstetricians. *M. = 16. **F. = 16 + x. 
Janssens, F. A. P. & Willems, J. 1908. 

Arachnida 

Epeira sclopetaria. M. = 11. F. = 11 + x. 
Berry, E. H. 1906. 

Coleoptera 

Anomoglossus emarginatus. M. = a. F. = a + x. 

Stevens, N. M, 1906. 
Chrysomela sirailis. M. = 11. F, = 11 + x. 

Stevens, N. M. 1909. 
Diabrotica. 12-punctata. M. = 9. F. = 9 + x. 

Stevens, N. M. 1908a. 
Diabrotica soror. M. = 9. F. - 9 + x. 

Stevens, N. M. 1908a. 
Diabrotica vittata. M. = 10. F. = 10 + x. 

Stevens, N, M. 1908a. 
Elater sp. M. = 9. F. = 9 + x. 

Stevens, N. M. 1906. 
Ellychnia corrusca. M. = 9. F. = 9 + x. 

Stevens, N. M. 1909. 
Hydrophilus piceus. M. = 15. F. = 15 + x. 

Arnold, G. 1909. 
Limoneus griseus. M. = 8. F. = 8 + x, 

Stevens, N. M. 1909. 
Necrophorus sayi. M. = 6. F. = 6 + x. 

Stevens, N. M. 1909. 
Phctinus consanguineus. M, = 9. F. = 9 + x. 

Stevens, N. M. 1909. 
Photinus pennsylvanicus. M. = 9. F. « 9 + x. 

Stevens, N. M. 1909. 
Stenopelmatus. M, = 23. F. = 23 + x. 
^ Stevens, N. M. 1909. 
M = male determining 
**F = female determining 



-3- 



Corrodentia 

Ceraatipeocue venosus. M. = 8. F. = 8 + x. 
Boring, A. M. 1913. 

Echinodermata 

Hipponde eeculenta. M. = a. F. = a + x, 

Tennent, D. H. 1911. 
Toxopneustes variegatue. M. = a. F. = a + x. 

Tennent, D. H. 1913. 

Hemiptera 

Agallia sanguinolenta. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Alydus eurinus. M. = 6, F. = 6 + x. 

Montgomery, T. H, 1906. 
Alydus piiosulus, M. = 6. F. = 6 + x. 

Wilson, E. B. 1905, b & c, 1906. 
Amphiscepa bivittata. M. = 13. F. = 13 + x. 

Boring, A. M. 1907. 
Anasa armigera. M. = 10. F. = 10 + x. 

Montgomery, T. H. 1906. 
Anasa sp. M. = 10. F. = 10 + x. 

Montgomery, T. H. 1906. 
Anasa tristis. M. = 10. F. = 10 + x. 

Wilson, E. B. 1905, b & c, 1906, 1907a. 
Aphrophora 4 notata. M. = 13. F. = 13 + x. 

Boring, A. M. 1907. 
Aphrophora quadrangular! s. M. = 11. F. = 11 + x. 

Stevens. N. M. 1906. 
Aphrophora spumaria. M, = 11. F. = 11 + x. 

Boring, A. M. 1907. 
Archimerus calcarator. M. = 7. F. = 7 + x. 

Wilson, E. B. 1905. 
Atymna castanea. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Campylenchia ourvata. M. = 9. F. = 9 + x. 

Boring, A. M. 1907. 
Catorintha. M. = 13. F. = 13 + x. 

Wilson, E. B. 1907. 
Ceresa bubalus. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Ceresa diceros. M. = 10, F. = 10 + x. 

Boring, A. M. 1907. 
Ceresa taurina. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Chariesterus antennator. M. = 13. F, = 13 + x. 

Wilson, E. B. 1909. 
Chelinidea. M. = 10. F. = 10 + x. 

Wilson, E. B. 1907. Morrill, C. V. 1910. 
Chlorotettrix unicolor. M. = 8. F. = 8 + x. 

Boring, A. M. 1907. 
Chlorotetrix vividus. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Cicada tibicen. M. = 13. F. = 13 + x. 

Wilcox. 1895. 



-4- 



Hemiptera 

Claetoptera obtusa. M. = 7. F. * 7 + x. 

Boring. A. M. 1907. 
Corizus alternatuB. M. = 6. F. = 6 + x. 

Montgomery, T. H. 1906. 
Corizus lateralus. M. = 6. F. = 6 + x. 

Montgomery, T. H. 1906. 
Corynoooris diatinctus. M. = 13. F. 13 + x. 

Wilson, E. B. 1909. 
Diedrocephala coccinea. M. = 11. F. * 11 + x. 

Boring, A. M. 1907. 
Diedrocephala mollipes. M. = 11. F. = 11 + x. 

Boring, A. M, 1907. 
Enchenopa binotata, M. = 9. F, = 9 + x. 

Boring, A. M. 1907. 
Entila sinuata. M. = 10. F. = 10 + x. 

Boring, A. M. 1907. 
Euthoctha galeator. M. = 10, F. = 10 + x. 

Wilson, E. B. 1907. 
Karmostes reflexulus. M. = 6. F. = 6 + x. 

Montgomery, T. H. 1901, 1906. 
Hygotrechus sp. M. * 10. F. = 10 + x. 

Montgomery, T. H. 1906. 
Largus cinctus. M. = 5. F. = 5 + x. 

Wilson, E. B. 1907, 1909, 1913. 
Largus sucoinctus. M. = 6. F. = 6 + x. 

Wilson, E, B. 1907. 
Leptocoris trivittatus. M. = 6. F. = 6 + x. 

Wilson, E. B. 1907. 
Leptoglossus phyllopus. M. = 10. F. = 10 + x. 

Wilson, E. B. 1907, 
Limotreohus marginatus. M. = 10. F. = 10 + x. 

Montgomery, T. H. 1906. 
Lygus pratensis. M. = 17. F. = 17 + x. 

Montgomery, T. H. 1906. 
Margus inconspicuus. M. = 11. F. = 11 + x. 

Wilson, E. B. 1907. 
Narnia. M. = 10. F. = 10 + x. 

Wilson, E. B. 1907. 
Oedencala dorsalis. M. = 6. F. = 6 + x. 

Montgomery, T. H, 1906. 
Pachylis gigas. M. = 7. F. = 7 + x. 

Wilson, E. B. 1907. 
Philoenus spumarius. M. - 11. F. = 11 + x. 

Boring, A. M. 1907. 
Phlepsius irrotatue. M. = 7. F. = 7 + x. 

Boring, A. M. 1907. 
Phymata sp. M, = 14. F. = 14 + x. 

Montgomery, T. H. 1906. 
Poeciloptera bivittata. M. = 13. F. = 13 + x. 

Boring, A. M. 1907. 
Poeciloptera pruinosa, M. = 13. F. = 13 + x. 

Boring, A. M. 1907. 



-5- 



Hemiptera 

Poeciloptera eeptentrionalis. M. = 13. F. = 13 + x. 

Boring, A. M. 1907. 
Protenor belfragi. M. = 6. F. = 6 + x. 

Montgomery, T, H. 1901, 1906. 
Pyrrochoris apteris. M. = 11. F. = 11 + x. 

Henking, H. 1891. Wilson, E. B. 1907. 
Vanduzea arcuata. M. = 8. F. = 8 + x. 

Boring, A. M. 1907. 

Myriapoda 

Lithobius ep. M. = a. F. = a + x, 

Elackman. 1907. 
Scolopendra heros. M. = 16. F. = 16 + x. 

Elackman, M. W. 1905, 1910. 
Scolopendra subspinipee. M. = 16. F. = 16 + x. 

Elackman, M. W. 1905, 1907. 
Scutigera forceps. M. = 18. F. = 18 + x. 

Medes, G. 1905. 

Nematoda 

Ancyracanthus cystidicola. M. = 5. F. = 5 + x. 

Mulson, K, 1912. 
Ascaris megalocsphala. M. = a. F. = a + x. 

Edwards. 1910. 
Creseis aclcula, M. = 9. F. = 9 + x. 

Zarnik, B, 1913. 
Heterakis dispar. M. = 4. F. = 4 + x. 

Gulick, A. 1911. 
Heterakis inflexa. M. = 4, F. = 4 + x. 

Gulick, A. 1911. 
Heterakis vesicularis. M. = 4. F. = 4 + x. 

Gulick, A. 1911. 
Strongylus paradoxus. M. = 5. F. = 5 + x. 

Gulick, A, 1911. 
Strongylus tenuis. M. = 5. F. = 5 + x. 

Gulick, A. 1911. 

Odonata 

Anax Junius. M. = 13. F. = 13 + x. 
Lefevre, G. & McGill, C. 1908. 

Orthoptera 

Anabrus sp. M. = 16. F. = 16 + x, 

McClung, C. E. 1903. 
Aplopus mayeri. M. = 17. F. = 17 + x. 

Jordan, H. E. 1908. 
Arphia simplex. M. = 11. F. = 11 + x. 

Carothers, E. 1913. 
Arphia tenebrosa. M. = 11. F. = 11 + x. 

Davis, H. 8. 1908. 
Blatta germanica. M. = 11. F. = 11 + x. 

Morse, M. 1909. 



-6- 



Orthoptera 

Blatta gerraanica. M. = 11. F. = 11 + x. 

Stevens, N. M. 1906, 
Brachyatola magna. M. = 11. F. = 11 + x. 

Sutton, W. S. 1900. 
Ceuthophilua ep. M. = 18. F. = 18 + x. 

Stevens, N. M. 1912, 
Chortophaga viridifasciata, M. = 11. F. = 11 + x. 

Davis, H. S. 1908, 
Chrysochjaon dispar. M. = a. F. = a + x. 

Veaely, J. 1913. 
Deoticus verrucosus. M. = 15. F. = 15 + x. 

Buchner, P. 1907. 
Diestraramena marmorata. M, = 28. F. = 28 + x. 

Schellenberg, A. 1913. 
Dissosteira Carolina. M. = 11. F. = 11 + x. 

Davis, H. S. 1908. Carothers, E. 1913. 
Gryllus assirailis. M. = 14. F. = 14 + x. 

Baumgartner, W. J. 
Gryllus campestris. M. = 19. F. = 19 + x. 

Buchner, P. 1909. 
Gryllus domesticus. M. = 10. F. = 10 + x. 

Baumgartner, W. J. 1904. 
Gryllotalpa borealis. M. = 11. F. = 11 + x. 

Payne, F. 1912, 
Gryllotalpa vulgaris. M, = 11. F. = 11 + x. 

Vom Rath. 1892. 
Hesperotettix prattensis. M. = 11, F. = 11 + x, 

McClung, C. E. 1905. 
Hesperotettix speciosus. M. = 11. F, = 11 + x, 

McClung, C, E. 1905. 
Hesperotettix viridis. M. = 11. F. = 11 + x. 

McClung, C. E. 1905. 
Hippiscus phoeniooptenis. M. « 11, F. = 11 + x. 

McClung, C. E. 1900. 
Hippiscus sp, M. = 11. F. = 11 + x. 

Buchner, P. 1909. 
Hippiscus tuberculatus. M. = 11, F. = 11 + x. 

Davis, H. S. 1908. 
Leptynia attenuata. M. = 17. F. = 17 + x, 

de Sinety. 1901. 
Leucophaea maderiae. M. = 11. F. = 11 + x, 

Morse, M. 1909. 
Locusta viridissima. M. = 16. F, = 16 + x. 

Otte, H. 1906. 
MelanopluB bivittatus. M, = 11, F. = 11 + x. 

Nowlin, N, 1908. 
Melanoplus feraoratus. M. = 11, F. = 11 + x. 

Davis, H. S. 1908. 
Melanoplus, femur rubrum. M. = 11. F. = 11 + x. 

Wilcox, E. V. 1895, 
Merrairia bivittata. M, = 11. F. = 11 + x. 

McClung, C. E. 1905. 



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Orthoptera 

MicDcentra ep. M. = 16. F. = 16 + x. 

McClung, C. E. 1908. 
Oedipoda. F. - 11. F. «= H + x. 

Buchner, P. 1909. 
Oedipoda miniata. M. = a. F. = a + x. 

de Sinety, R. 1901. 
Orcheeticus sp. M, = 16. F. = 16 + x. 

McClung, C. E. 1902. 
Orphania denticauda. M. = 15. F. = 15 + x. 

de Sinety, R. 1901. 
Psunphagus marmoratus. M. = 9. F. = 9 + x. 

Giglio, Tos. 1908. 
Periplanata americana. M. = 11. F. = 11 + x. 

Morse, M. 1909. 
Pezotettix. M. = 11. F. = 11 + x. 

Buchner, P. 1909. 
Phrynotettix raagnus. M. = 11, F. = 11 + x. 

Pinney, E. 1908. 
Psophua. M. = 11. F. = 11 + x. 

Buchner, P. 1909. 
Schistocerca alutacca. M. = 11. F. = 11 + x. 

Hartman, F. 1913. 
Schistocerca americana. M. = 11. F. = 11 + x. 

Hartman, F. 1913a. 
Scudderia sp. M. = 16. F. = 16 + x. 

McClung, C. E. 1902. 
Steiroxys trilineata. M. = 14. F. = 14 + x. 

Davis, H. S. 1908. 
Stenobothrus biguttulus. M. = 8. F. = 8 + x. 

Gerard, Pol. 1909. 
Stenobothrus curtipennis. M. = 8. F. = 8 + x. 

Davis, H. S. 1908. 
Stenobothrus parallelua. M. = 8. F. = 8 + x. 

de Sinety, R. 1901. 
Stenobothrus viridulus. M. = a. F. = a + x. 

Meek, C. F. U. 1911 & 1912. 
Stenopelmatus. M. = 23. F. = 23 + x. 

Stevens, N. M. 1905, 
Stylopyga orientalis. M. = 11. F. = 11 + x. 

Morse, M. 1909. 
Syrbula acuticornis. M. = 11. F. = 11 + x. 

Robertson. 1908. 
Syrbula admirabilis. M. = 11. F. = 11 + x. 

Robertson, W. R. B. 1908. 
Syrbula fusca vittata. M. = 11. F. = 11 + x. 

Robertson, W. R. B. 1908. 
Xiphidiura fasciat\im. M. = 16. F. = 16 + x. 

McClung, C. E. 1902. 



Vertebrata 

Didelphys virginiana (opossum). M. = 8. F. = 8 + x 
Jordan, H. E. 1911. 



-8- 



TYPE II. n/2 + y, n/S + x 
Annelida 

Tomopteris oniscif ormis. M. = 8 + y. F. = 8 + x. 

Shreiner, A. & K. E. 1906. 
Zoogonus mirus. M, = 4 + y. F. = 4 + x. 

Goldschmidt, R. 1905. 

Araohnida 

Lyoosa insopita. M. » 12 + y. F. = 13 + x. 
Montgomery, T. H. 1905. 

Coleoptera 

Adalia blpunctata. M. = 9 + y. F. = 9 + x. 

Stevens, N. M. 1909. 
Blepharida rhois. M. = 15 + y. F. = 15 + x. 

Stevens, N. M. 1906. 
Buprestidae. M. = a + y. F. = a + x. 

Stevens, N. M. 1906. 
Chelymorpha orgus. M. = 10 + y. F. = 10 + x. 

Stevens, N. M. 1905, 
Chlaemus aestivus. M. = a + y. F. = a + x. 

Stevens, N. M. 1906. 
Chlaenius pennsylvanicus. M. = a + y. F. = a + x, 

Stevens, N. M, 1906. 
Chrysochus auratus. M. = 12 + y. F. = 12 + x, 

Stevens, N. M. 1909, 
Cicindela primeriana. M. = 9 + y, F. = 9 + x, 

Stevens, N. M. 1906, 
Cicindela vulgaris. M. = 10 + y. F, = 10 + x. 

Stevens, N. M, 1909. 
Coptocyia aurichalcea. M. = 10 + y. F. = 10 + x. 

iNfowlin, N. W. 1906. 
Coptocycla clavata. M. = 8 + y. F. = 8 + x. 

Stevens, N. M. 1909. 
Coptocvcla guttata. H. = 8 + y, F. = 8 + x. 

Nowlin, N. W. 1906, 
Cylene robinia. M. = 9 + y...F, = 9 + x. 

Stevens, N. M. 1909, 
Doryphora clivicolis. M. = 16 + y. F. = 16 + x, 

Stevens, N. M, 1909, 
Doryphoria deoemlineata. M. = 17 + y. F. « 17 + 

Stevens, N. M. 1906. 
Epicauta cinerea. M. = 9 + y. F. = 9 + x. 

Stevens, N. M. 1909, 
Epicauta pennsylvanica. M. = 9 + y. F. = 9 + x, 

Stevens, N, M. 1909. 
Epilachna borealis. M, = 7 + y. F. = 7 + x. 

Stevens, N. M. 1906. 
Euchroma gigantica. M. 12 + y. F. = 12 + x. 

Nichols, M. L, 1910, 
Euphoria inda. M. = 9 + y, F. = 9 + x, 

Stevens, N. M. 1906. 



-9- 



Coleoptera 

Galerita bicolor. M. = a + y. F. = a + x. 

Stevens, N. M. 1906. 
Haltica chalybea. M. = 10 + y. F. = 10 + x. 

Stevens, N. M. 1909. 
Lema trilineata. M. = 15 + y. F. = 15 + x. 

Stevens, N. M. 1909. 
I.ina laponlca, M. = 16 + y. F. = 16 + x. 

Stevens, N. M. 1909. 
liistotrophus cingulatus. M. « 12 + y. F. = 13 + x 

Stevens, N. M. 1909. 
Obera tripunctata. M. = a + y. F. = a + x, 

Stevens, N. M. 1909. 
Odontota dorsalis. M. = 7 + y. F. = 7 + x. 

Stevens, N. M. 1906. 
Penthe obliquata. M. = 7 + y. F. = 7 + x. 

Stevens, N. M. 1909. 
Phytonomius punctata. M. = a + y. F. = a + x. 

Stevens, N. M. 1909. 
Silpha americana. M. = 19 + y. F. = 19 + x, 

Stevens, N. M. 1906. 
Staphylinus violaceus. M. = 21 + y. F. « 21 + x. 

Stevens, N. M. 1909. 
Tenebrio molitor. M. = 9 + y. F. = 9 + x. 

Stevens, N. M. 1905. 
Tetraopes tetraophthalmus. M. = 9 + y. F. = 9 + x 

Stevens, N. M. 1909. 
Trirhabda oanadense. M. « 14 + y. F. = 14 + x. 

Stevens, N. M. 1906. 
Trirhabda virgata. M. = 13 + y. F. = 13 + x. 

Stevens, N. M. 1906. 

Diptera 

Anapheles pinctpennis. M, - 5 + y. F. = 5 + x. 

Stevens, N. M. 1910. 
Calliphora vomitoria. M. = 5 + y. F. - 5 + x. 

Stevens, N. M. 1908a. 
Culex pipiens. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1910. 
Culex tarsilis. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1910. 
Drosophila amoena. M. « 3 + y. F. - 3 + x. 

Metz, C. W. 1914. 
Drosophila ampelophila. M. = 3 + y. F. = 3 + x. 

Stevens, N. M. 1908a. 
Drosophila funebris. 1. - 4 + y. F. = 4 + x. 

Metz, C. W. 1914. 
Drosophila, quinaria. M. = 3 + y. F. = 3 + x. 

Metz, C. W. 1914. 
Drosophila repleta. M. = 5 + y. F. = 5 + x. 

Metz, C. W. 1914. 
Eristalis tenax. M. = 5 + y. F. = 5 + x, 

Stevens, N. M. 1908a. 



-10- 



Diptera 

Lucilia Caeaar. M. = a + y. F. = a + x. 

Stevens, N. M. 1908. 
Musoa domestica. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1908. 
Phorbia brassica. M. = a + y. F. = a + x. 

Stevens, N. M. 1908. 
Sarcophaga sarraciniae. M. = 5 + y. F. = 5 + x 

Stevens, N. M. 190Sa. 
Scatophaga pallida. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1908a. 
Tetanocera sparsa. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1908a. 
Theobaldia incidens. M. = 5 + y. F. = 5 + x. 

Stevens, N. M. 1910. 

Hemiptera 

Apiomerls crassipes. M. = 11 + y. F. = 11 + x. 

Payne, F. 191S. 
Banasa calva. M. = 13 + y. F. = 13 + x. 

inison, E. B. 1907b. 
Banasa diraidiata. M. = 7 + y. F. = 7 + x. 

Wilson, E. B. 1907b. 
Brochymena. M. « 6 + y. F. = 6 + x. 

Wilson, E. B. 
Calocoris rapidus. M. = 14 + y. F. = 14 + x. 

Montgomery, T. H. 1906. 
Coenus delius. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1905. 
Cosmopepla carnifex. M. « 7 + y. F. = 7 + x. 

Montgomery, T. H. 1906. 
Diplocodus exsanguis. M. = 13 + y. F. = 13 + x 

Payne, F^ 1909. 
Enchenopa binotata. M. = 9 + y. F. = 9 x. 

Kornhauser, S. I. 1914. 
Eurygaster alternatus. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Euschistus crassus. M. = 5 + y. F. = 5 + x. 

Foot & Strobell. 1913. 
Euschistus fissilis. M. = 6 + y. F. = 6 + x. 

Wilson, E. E. 1905. (b & c) 1906. 
Euschistus ictericus. M. = 6 + y. F. = 6 + x. 

Wilson, E. E. 1906. 
Euschistus servus. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1905. 
Euschistus tristigmus. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1901, 1906, 
Euschistus variolarius. M. = 6 + y. F. = 6 + x 

Wilson, E. B. 1906. 
Lygaeus bicruris. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1913. 
Lygaeus turcicus. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1905. (a & b) 1906. 



-11- 



Hemiptera 



Metapodius (acanthocephala) feraoratue. /M. = 10 + y. 

Wilson, E. E. 1909. IF. = 10 + x. 

Metapodius (acanthocephala) granulosus. |M. = 10 + y. 

Wilson, E. B. 1909. \f. = 10 + x. 

Metapodius (acanthocephala) terminalis. fM, = 10 ■+ y. 

Wilson, E. B. 1909. (F. = 10 + x. 

Mormidea lugens. M. = 6 + y. F. = 6 + y, 

Montgomery, T. H. 1906. 
Nabis annulatus. M. = 8 + y. F. = 8 + x. 

Montgomery, T. H. 1906. 
Nezara Hilaris. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1905. Wilson, E. B. 1910. 
Nezara viridula. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1910. 
Oebalus pugnax. M. = 4 + y. F. = 4 + x. 

Wilson, E. B. 1910. 
Oncopeltus fasciatus. M. = 7 + y. F. = 7 + x. 

Wilson, E. B. 1913. 
Peliopelta abbreviata. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Peribalus lirabolaris. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Perillus confluens. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Podissus spinosus, M. = 7 + y. F. = 7 + x. 
Montgomery, T. H. 1901, 1906. 
Wilson, E. B. 1906, 
Poecilocapsus goniphorus. M. = 16 + y. F. = 16 + x. 

Montgomery, T. H. 1906. 
Rocconota annulicornis. M. = IS + y. F. = 13 + 3x. 

Payne, F. 1909. 
Stiretrus anchorago. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1909. 
Tingis clavata. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Trichopepla seraivittata. M. = 6 + y. F. = 6 + x. 

Montgomery, T. H. 1906. 
Trichopepla. M. = 6 + y. F. = 6 + x. 

Wilson, E. B. 1905. 
Zaitha sp. M. = 11 + y. F. = 11 + x. 
Montgomery, T. H. 1906. 

Nematoda 

Ascaris felis. M. = 8 + y. F. = 8 + x. 
Edwards. 1911. 



Orthoptera. 

Anisolabis maritima. M. = 11 + y. F. = 11 + x. 

Randolph, H. 1908. 
Forficula auricularia. M. = 11 + y. F. = 11 + x. 

Stevens, N. M. 1910. 

de Sinety. 1901. 



12- 



TYPE III. n/2, n/2 + 2 or more x. 
Arachnida 

Agalena naevia. M. = 19. F. = 19 + 2x. 

Wallace, L. B. 1908. 
Pholcus phalange idee. M. = a. F. = a + 2x. 

Wallace, L. B. 1908, 
Maevia vittata. M. = a. F. = a + 2x. 

Painter, T. S. 1913. 

Hemipt era 

Syromastes marginatus. M. = 10. F. = 10 + 2x. 
Qross, J. 1904. 

Nematoda 

Ascarie lurabricoidee. M. = 19, F. = 19 + 5x. 
Edwards. 1910. 

Trenatoda 

Schistosomum haematobium. M. = 6. F. = 6 + 2x. 
Lindner, E. 1914. 

Vertebrata 

Pig. M. = 10. F. 10 + 2x. 

Wodsedalek, J. E. 1913. 
Homo sapiens. (man) M. = 10. F. = 10 + 2xf 

Guyer, M. F. 1910. 

Winiwarter, H. von. 1912. M. = 23. F. = 24 



TYPE IV. n/2 + y, n/2 + 2 or more x. 
Hemiptera 

Acholla ampliata. M. = 10 + y. F. = 10 + 5x. 

Payne, F. 1909. 
Conorhinue sanguisugus. M. = 10 + y. F. = 10 + 2 

Payne, F. 1909. 
Fitchia spinosula. M. = 13 + y. F. = 12 + 2x. 

Payne, F. 1909. 
Gelastocoris (Galgulue) oculatus. M. = 15 + y. F 

Payne, F. 1909. 
Pnirontie modesta. M. = 10 + y. F. = 10 + 4x. 

Payne, F. 1912. 
Peelliodes cinctus. M. = 12 + y. F. = 12 + 3x. 

Payne, F. 1912. 
Prionidue cristatus. M. = 11 + y. F. = 11 + 3x. 

Payne, F. 1909. 
Thyanta calceata. M. = a + y. F. = a + 2x. 

Wilson, E. B. 1909. 
Sinea complexa. M. = 12 + y. F. = 12 + 3x. 

Payne, F. 1912. 
Sinea confusa. M. = 12 + y. F. = 12 + 3x. 

Payne, F. 1912. 



-13- 



Heraiptera 

Sinea diadema. M. = 12 + y. F. = 12 + 3x. 
Payne, F. 1909, 1912. 

Hemiptera 

Sinea rileyi. M. = 12 + y. F. = 12 + 5x. 

Payne, F. 1912. 
Sinea spinipee. M. = 12 + y. F. = 12 + 3x. 

Payne, F. 1912. 

Nematoda 

Ascaris lurrbricoidee. M. = 18 + x. F. = 18 + 5x. 
Edwards, C. L. 1910. 



TYPE V. n/2, n/2. (No observable difference in spermatozoa. 
There may or may not be a difference In the eggs. ) 

Amphibia 

Salamandra maculosa. M. = 12. F. = 12. 
Schreiner, A. & Z. E. 1906b, 

Coleoptera 

Silpha carinata. M. =16. F. =16. 
Holmgren, N. 1902. 

Echinodermata 

Aeterias vulgaris. M. =9. F. = 9. 
Tennent, D. H. 1907. 

Hemiptera 

Brown Rose aphid. M. =5. F. = 5. 
Green Rose aphid. M. =7. F. = 7. 
Migratory Rose aphid. M. =9. F. = 9. 
Saranac Willow aphid. M. =5. F. = 5, 
Harpswell Willow aphid. M. = 3. F. = 3. 
Aphis oenotherae. M. =5. F. = 5, 
Oenothera aphid II. M. =4. F. = 4. 
Black milkweed aphid. M. =4. F. = 4. 
Orange milkweed aphid. M. =4. F. = 4. 
Pale milkweed aphid. M. =7. F. = 7. 
Nasturtium aphid. M. = ^. F. = 4. 
Oak aphid. M. = 7. F. = 7. 
Beach goldenrod aphid. M, =6. F. = 6. 
Tall goldenrod aphid. M. =6. F. = 6. 
Paper birch aphid. M. = 9. F. = 9. 
Clover aphid. M. =8. F. = 8. 
Woolly beech aphid. M. = 8..F. =8. 
Star cucumber aphid. M. =5, F. = 5. 
Maple aphid. M. = a. F. = a. 
Pea aphid. M. =4. F. = 4. 
Gourni aphid. M. =5. F, = 5. 
Stevens, N. M. 1905. 



-14- 



leoptera 

TermopeiB angusticollis. M. 26. 
Stevena, N. M. 1905. 

Lepldoptera 

Abraxas. M. = 9. F, =■• 9. 

Wilson, E. B. 1909. 
Cacoecia cerasivorana. M. = a. F 

Stevens, N. M. 1906. 
Euvanessa antiopa. M. = a. F. = 

Stevens, N. M. 1906, 

Mollusca 

Sagitta bipunctata. M. =9. F. = 
Bordas, M. 1913. 

Trichoptera 

Platyphylax designatus. M, = 30. 
Lutman, B. F. 1910. 



UNCLASSIFIED FORMS HAVING SEX CHROMOSOMES: 

Vertebrata 

Daeypus sexcinctus (armadillo) 
Patterson, N. F. 1910. 

Bat. 

Jordan, H. E. 1913. 
Bos tauris (bull) 

Jordan, H. E. 1913. 

Scbnoenfeld, H, 1901. 
Felie catus. (cat) 

Saintmont, J, 1909. 
Ca^nis farailiaris (dog) 

Jordan, H. E. 191:^. 
Equus caballus (horse) 

Jordan, H. E. 1913. 
Guinea pig. 

Stevens, N. M. 1911. 

Mule. 

Jordan, H. E. 1913. 

Rat. 

Duesberg, J. 1909. 
Ovis (sheep). 

Jordan, H. E. 1913. 
Mus sp. (white mouse) 

Jordan, H. E. 1913. 



-15- 

4« Prevl ouB Work on Siz e Dlmorp hlBm 
Dr. Charles Zeleny and Mr. K. C. Fauat published data 
and curves for fifteen species of animals in the Journal of Ex- 
perimental Zoology, Volume 18, Number 2, February, 1915. Alto- 
gether they made thirty-three separate determinations. The sub- 
jects used were Musca domeetica, Lygaeus kalmii, Alydus piloaulus, 
Anasa tristis, Trirhabda toraentosa, Phytonomus punctatus, Mela- 
noplue femur rubrum, Melanoplus dif f erentialis, Gryllus abbrevia- 
tus, Aeshna canadensis, Rana pipiens, Pseudemys troosti, Ovis aries, 
Eos taurus and Canis familiaris. Their general results indicate 
that the population of spermatozoa from a single testes is made 
up of two separate groups. 

Mr. J. E. Wodsedalek published data on the pig in 1913 
which indicates a bimodal curve in the spermatozoan population of 
this animal. 

II. MATERIAL AND METHODS 
1. Most Desirable Speci es 
The best species which could be used in a piece of work 
of this character would be those in v*rhich the greatest chromosom- 
al differences have been described. Such forms as Protenor bel- 
fragei, Largu.s cinctus and others of like nature seemed highly 
desirable. Forms for which figures of the chromosomes were pub- 
lished were also desirable, as a calculated ratio obtained from 
these figures would offer a standard for checking the actual ratio 
obtained. The lengths of tte spermatozoa should vary as the cube 
root of the total volume of the contained chromatin. 



-16- 

The most desira-ble species were not easily obtainable 
inasmuch as in many cases sexual maturity occurs during the sum- 
mer. Material from such species as could be found was mounted 
and that which was seemingly the most desirable was used in the 
measurements. A considerable number of insects hibernate be- 
neath old boards, decaying bark, etc. The bases of mullein plants 
often harbor many guests when other places are barren of life. 

Insect testes are always located in the abdomen, usually 
towards the tip and at the dorsal side. They are readily dis- 
tinguished and are often yellow or red in color. 

2. Fixat ion and Staini ng: 

After experimenting with Delafield, Ehrlich and Heiden- 
hain haeraatoxylins it was finally decided that Heidenhain's iron 
haenatoxylin gave the best results. The testes were dissected 
in normal salt solution. The spermatozoa, taken when possible 
from the vas deferens, were smeared on an albumen coated slide 
and fixed over osraic acid fumes. After a few seconds of treatment 
with the osraic fumes they were allowed to just reach a dry condi- 
tion, then washed in tap water and left in Heidenhain for several 
hours. From the Heidenhain they were dipped in tap water and 
left in a one half per cent solution of haematoxylin for 10-15 
hours. The time factor seems to be unimportant if not too serious 
ly abused. They were rinsed in tap water again and destained in 
the Heidenhain. Destaining takes about eight minutes with a fresh 
solution. When properly destained they were run up thru the al- 
cohol series into xylol and mounted in balsam. In bringing the 



-17- 

elides over into the one-half per cent haematoxylin, a slight 
amount of the Heidenhain is carried also. The iron in this aids 
in giving a desirable intensely black stain. 

3. Method of Measur ement 
In all the sperm heads measured, the natural form seemed 
to be a straight one. Many individuals were curved and twisted 
on the slide. Of course, only the straight ones were measured. 
It is necessary to accuracy that the spermatozoan be in a horizon- 
tal position. No individuals were measured, all of whose length 
was not visible at the same amount of focus in the microscope. 
Measurements were made with a No. 2 Leitz ocular, A l/lZ oil 
immersion objective was used. A mechanical stage obviated all 
danger of duplicate measurements. In practically all cases sperma- 
tozoa were measured to lengths of an ocular division. Personal 
errors were eliminated as far as possible by numerous remeasure- 
ments. Measurements were not made by poor light. No measurements 
were made when fatigue might influence the result. Care was 
taken to have all measurements made under as nearly the same con- 
ditions as possible. Tabulation of data was not made until the 
full number decided upon for the set had been completed. This 
was to avoid danger of a false judgment and the selection of par- 
ticular values. 

4. Source s of Possible Error 
Before accepting the results obtained all possibilities 
of incorrectness must be considered. These fall into three 
classes: 1. Mechanical erroB, those due to the personal equation 
of the observer or other errors of measurement. 3. Errors due 



-18- 



to poor methods of technique. 3. Errors due to artificial demor- 
phiam in the living material which might ensue because of the 
presence of any one of a number of conditions. 

k» Possible Mechanical Errors 

a. All spermatozoa, all of whose length was not in 
focus at the same time, were eliminated. It is possible that this 
might occur more frequently with long individuals than with short 
ones. This might produce asymmetry in a curve but there is no 
reason to believe that it would result in bimodality. 

b. Each set of measurements required a number of sit- 
tings, usually twelve to fifteen. This number itself would tend 
to prevent bimodality resulting from possible change of standards, 
poor light, fatigue, etc, 

0. The character of the scale might tend to cause 
grouping around certain points. Thick lines might obscure the 
nearest tenths, etc. The scale used had the narrowest lines of 
any obtainable. Frequent remeasurements were made and final re- 
sults did not show any grouping of this kind. Many spermatozoa com« 
nearly between two units of measurement. Biased judgment resulting 
from previous determinations, or an unbased hypothesis might 
throw these into one group more frequently than another. In plot- 
ting curves the obtained lengths were grouped by fifths while 
measurements were made to tenths, thus counteracting this tendency. 

B. Possible Errors of Technique 
a. Lack of uniformity of fixation or staining on the 
slide. Tabulations made by Zeleny and Faust for different regions 



-19- 



of the slide ehowed no deviation from the general diraorphlem. 

b. Material at the edge dries more rapidly in the pro- 
cess of fixation than that of the center. Tabulations by Zeleny 
and Faust of material of the two regions showed no essential 
difference between them. 

o. Material drawn over a dlide by means of currents 
set up in the suspending liquid may have smaller individuals at 
one end of the smear than at the other. In the preparations made, 
all material was equally distributed to all parts of the slide and 
dried in position before staining. 

d. In smearing material some spermatozoa may be 
stretched or otherwise distorted. These are easily recognized 
and were always eliminated. 

e. Some spermatozoa may be curved. These were never 
measured, only the straight individuals being used. 

f. In case longer spermatozoa curved more often than 
shorter ones a disparity of numbers might occur. However, this 
would cause asymmetry rather than bimodality, 

g. The chromatin rod usually takes an intense stain 
and the cytoplasmic cap a light one. In cases of a short cap or 
insufficient destaining there might be a tendency to measure to 
the end of the head. This would produce apparent dimorphism. 

C. Errors Due to Artificial Dimorphism in the Living Material 
a. Error of random sampling. The spermatozoa taken 
may not be representative of the mass for the entire testis. The 
large number of individuals measured should prevent this possibilit] 



-20- 

"b. Two or more different degreeo of maturity may be 
preeent in the material studied. Thia would give an apparent 
dimorphiem. The use of material which was very active would tend 
to eliminate this chance. The supposition would also involve 
the hypothesis that there was an abrupt bree^k in continuity be- 
tween two stages of development. This seems to be hardly likely, 

o. Spermatozoa of like size may congreg-ate in masses 
in particular parts of the vae deferens because of similarity in 
physiological activity or perhaps mechanically. The entire con- 
tents of the vas deferens were suspended in a drop of physiological 
normal salt solution and the whole was thoroughly mixed before 
making smears. 

d. The chromatin rod length may not be a true index of 
the length of the entire sperm head. As the chromatin rod repre- 
sents nearly the entire length this objection is not to be con- 
sidered seriously. Moreover quantitative difference in chromatin 
content ciuet necessarily affect the head length of spermatozoa. 

e. Spermatozoa from two different individuals might 
show dimorphism not due to differences present within a single 
testes. The fact that material used was taken from a single 
testis removes this difficulty. 

Other sources of error might be included. Tho se cited 
are the ones regarded as most important. T^rea-rly all these possi- 
bilities would cause asymmetry rather than bimodality. Moreover 
the accumulation of data, practically all of which shows bimodality, 
makes the supposition nearly incontrovertible that such bimodality 
is due to the presence of two kinds of spermatozoa differing in 



a 



c 



d 



f 



i 



h 



Spermatozoan heads 

a. Leptocorls trivittatua 

Id. Reduviolus ferus 

0. Corizus lateralus 

d. Euschistus variolariue 

e. Cosmopepla carnifex 

f. Passalus cornutus 

g. Berosus striatus 

h. HeliodriluB caliginosa 

Magnification = 900 



OF THE 
UNIVERSITV OF ILLINOIS 



-21- 



chroniatin content. 

DATA 
T epto corle triyittatue 

E. B, Wil6on (1906) states that this insect has two 
kinds of spermatids, one containing seven and the other six chromo- 
somes. This would put it in Type 1. No figures could "be obtained. 
A rough comparison can be made by using the expected ratio of 
Anasa tristis, another coreid, as its measure. 

Material was obtained at Urbana in early March. One 
testis, only, was used and this gave an abundance of straight 
active spermatozoa. Nine hundred and eighty four measurements were 
made. One unit on the ocular micrometer scale equals 1.717 microns 
The resulting curve is noticeably bimodal, the modes coming at 
25.4 microns and 27.8 microns. The length ratio is 25.4:27.8 = 
1:1.09. The first five hundred measurements were used as a basis 
for a curve and the remainder for another curve. The results 
were bimodal curves with modes at 25.4 and 27.8. The expected 
ratio of Anasa tristis is 1:1.11. 

2. Reduviolus f eru s 

No cytological data could be obtained for this form 
but members of the same family have two kinds of spermatids accor- 
ding to T. H. Montgomery (1909). 

Material was collected at Urbana about the middle of 
March. An abundance of active spermatozoa was taken from a single 
testis. This gave plenty of straight heads after fixing and 
staining. Five hundred measurements were made. One unit on the 



LEPTOCORIS TRIVITTATUS 




1. Value in microns 30.9 31.3 21,6 33.0 23.3 23.7 33.0 23.3 



Frequency 



icy 




1 


2 


3 


6 


9 


13 


18 


23.7 


24.0 


24.4 


24.7 


25.0 


25.4 


25.7 


26.1 


26.4 


25 


26 


28 


30 


41 


53 


37 


31 


38 


27.1 


27.4 


27.8 


28.1 


28.4 


28.8 


29.1 


29, 5 


29,8 


41 


50 


60 


51 


45 


41 


39 


38 


40 


30. 5 


30.9 


31.2 


31.6 


31.9 


32.3 


32.6 


33 




24 


20 


15 


12 


11 


8 


7 


5 





23 



40 



31 



OF THE 

UNIVERSITY OF ILLlNOlb 



-22- 



ocular micrometer scale equals 1.717 microns. The curve indicates 
"bimodali ty, but would probably be improved by the addition of a 
large number of measurements. The modes occur at 27.4 microns, 
and 28.8 microns. The head length ratio is 27.4:28.8 = 1.00:1.05. 

3. CorizuB late raluB 

Montgomery (1906) states that this insect has two kinds 
of spermatids, one having six and the other seven chromosomes. 
This would place it in Type I. However, his figures are not large 
enough to be of value in determining the approximate chromatin 
ratio for the two spermatozoa. The expected ratio used was that 
of Anasa tristis, another coreid. 

Material was obtained at Urbana in late March and gave 
an abundance of lively spermatozoa. These fixed in a satisfactory 
manner. Two measurements, each of five hundred individuals, were 
made. One division of the ooular micrometer equals 1.717 microns. 
The second series is a check upon the first. In both cases bi- 
modal curves resulted. The modes were alike coming at 27.1 microns 
and 2S.5 microns. The head length ratio is 27.1:2S.5 = 1:1.09. 
The expected ratio, that of Anasa tristis is 1:1.11. This is 
probably a trifle high as Anasa tristis has an exceptionally large 
"x" chromosome. 

^* Euschi stus vari olarius 
Wilson (1906) demonstrates that there are two kinds of 
spermatids. The male producing ones have six normal chromosomes 
plus a small "y" shromosome a-nd the female producers have six 
normal chromosomes plus an "x" chromosome. This would place the 



I I j 

! 

REDUVIOLUS FERUS 4 




Value in microns 


33.7 


34.0 


34.4 


34.7 


35,0 


35,4 


35.7 


Frequency 




4 


5 


6 


5 


13 


19 


35 


36.1 


36,4 


36.7 


37.1 


37.4 


37.8 


38,1 


38.4 


38.8 


30 


35 


36 


49 


56 


45 


33 


37 


35 


39.1 


39.5 


39.8 


30.1 


30.5 


30.9 


31,3 


31.6 


31.9 


33 


18 


18 


10 


9 


8 


7 


5 


4 



33.3 33.6 
3 1 



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U. OF I. 6. S. FORM S 



3, Value in raicrona I. 
Frequency 



25.0 


25. 


4 


25.7 


13 


14 




16 


28.4 


28.8 


29.1 


13 


19 




26 


31.9 


32. 


3 


32.6 


6 


9 




5 








II. 


25.4 


25. 


7 


26.1 


11 


11 




13 


28.8 


29. 


1 


29. 5 


19 


28 




45 



23.0 


23.3 


23.7 


24.0 


24.4 


24. 


7 




7 


8 


10 


13 


13 


13 






26.1 


26.4 


36.7 


27.1 


27.4 


27. 


8 


28. 


19 


23 


27 


42 


30 


15 




11 


29. 5 


29.8 


30.1 


30.5 


30.9 


31. 


3 


31. 


38 


28 


20 


17 


11 


10 




6 


23.0 


23,3 


33.7 


24.0 


24.4 


24.7 


25! 


4 


6 


7 


7 


5 


7 




11 


26.4 


26.7 


37.1 


27.4 


27.8 


28. 


1 


28. 


20 


27 


54 


32 


20 


21 




17 


29.8 


30.1 


30.5 


30.9 


31.2 


31. 


6 


31. 


30 


15 


12 


10 


7 


8 




4 



32.3 
5 



32.5 
5 




U. OF I. S. S. rORM » 



OF THE 

UNIVERSITY OF ILLINOIS 



-23- 



Ineect in Type II. Wilson givee figures of the chromosomeB of 
this species on page 15 of the Journal of Experimental Zoology for 
1906. By computing their columee as cylinders the sum of the 
volumes of the chomosomee of the two kinds of spermatozoa may be 
obtained. The ratio of the cube root of these sums will be the 
expected ratio of the spermatozoan heads. Length varies as the 
cube root of volume. 



Chromosome 


Width 


Length 


Volume 


a 


2.8 


3.1 


19.085 


b 


2.5 


3.7 


18.16 


c 


2.6 


4.0 


21.24 


d 


2.8 


4.3 


26.48 


e 


2.2 


3.0 


11.41 


f 


1.9 


2.1 


5.95 


X 


1 


1 


0.79 


X 


2.4 


2.4 


10.86 



Chromatin volume of male producing spermatozoan = 103.1104. 
Chromatin volume of female producing spermatozoan = 113.1824. 
Calculated ratio = 1.00:1.04. 

Material was obtained in Urbana about the middle of 
April. There was evidence that all of the spermatozoa were not 
fully developed. This necessitated careful selection in making 
measurements so as not to include unripe ones. These latter are 
distingiaishable by their thickness and less definite outlines. 
Five hundred measurements were made. One division of the ocular 
micrometer equals 1.717 microns. The resulting curve is 




Value in raicrone 


13.0 


13.4 


13.7 


14.1 


14.4 


14.7 


15.1 


Frequency 


6 


16 


20 


21 


24 


46 


60 


15.5 15.8 


16.1 


16.5 


16.8 


17.2 


17.5 


17.9 


18.2 


39 38 


58 


84 


38 


12 


7 


6 


6 



18.5 
3 



1 



ttrr 



U. OF I. S. 



S. FORM 3 



OF THE 
UNIVERSITY OF ILLINOIS 



-24- 

conspicuously bimodal. The modes come at 15.1 microns and 16.5 
microns. The head langth ratio is 15.1:16,5 = 1:1.09 

5. Coamo pepla carn if ex 
T. H. Montgomery (1906) states that this species has two 
kinds of spermatids. The male producers have seven normal chromo- 
somes plus a "y" chromosome and the female producers have seven 
normal chromosomes plus an "x" chromosome. Thfe places the insect 
in Type II. Such figures as are given were useless for computing 
the probable head length ratio of the spermatozoa. This being 
a Pentatomid, the same expected ratio was taken for ittliatwas used 
for Euschistus variolarius. Material was obtained in Urbana in 
early May. The mass of spermatozoa was uniformly active and took 
a good fixation. Five hundred measurements were made. One divi- 
sion of the ocular micrometer equals 1. 71 7microns. The resulting 
curve shows well marked bimodality. The modes come at 18.5microns 
and 19.9 microns. The head length ratio is 18.5:19.9 = 1:1.075. 
The expected ratio is 1:1.05. 

6. Passa lus cornutus 
Ho published data could be obtained for this form. 
Many varieties of beetles have been described by N. M. Stevens 
and others. All. of these have "sex" chromosomes of one or 
another of the described types; Nos. I and II predominating. 

Material was obtained at Urbana in late March. The 
spermatozoa were uniformly active and made a very good preparation. 
Five hundred measurements were made. One division of the ocular 
micrometer scale equals 1.717 microns. The resulting curve tends 
towards unimodality. The slight projection at 12.4 may indicate 




5. Value in microns 


16.5 


16.8 


17.2 


17.5 


17.9 


18.2 


18.5 


Frequency 


6 


14 


16 


19 


26 


42 


83 


18.9 19.2 


19.6 


19.9 


20.3 


20,6 


20.9 


21,3 


21.6 


53 35 


42 


68 


37 


14 


11 


11 


9 



22.0 
8 







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U. OF t. 6. S. FORM 3 



UNlV£RSlTy^o"/lLUNa,. 



-25- 

either asymmetry or blmodality. A definite mode ocoure at 11.7 
microns. 

7, Berosus stria tue 
No published data is available for use in judging this 

form. 

Material was obtained at Urbana in early April and gave 
bounteously of active spermatozoa. Straight spermheads were in 
profusion. Five hundred measurements were made. One division of 
the ocular micrometer scale equals 1.717 microns. The resulting 
curve is distinctly biraodal. The modes occur at 16.1 microns 
and 17. S, The sperm head length ratio is then 15.1:17.2 = 1.00: 
1.07. 

8. Heliodr ilus caligin osa 

This species is very interesting as it is hermaphroditic. 
No doubt many zoologists would say without hesitation that a curve 
of the head lengths of its spermatozoa would be unimodal. Un- 
fortunately no published data giving a clear account of the sperma- 
togenesis of the earth worm could be obtained. 

Material was obtained at Urbana in April. Plenty of 
functional spermatozoa were taken from the sperm ducts. The pre- 
pared slides were not of the best quality as the sperm head out- 
lines were not uniformly distinct and tended to distortion. Only 
the best ones were measured. Five hundred measurements were made. 
One division of the ocular micrometer equals 1.717 microns. The 
plotted curve is flat topped. This would seem indicate bimodality, 
but may be due to an insufficient number of measurements. 



FASSALUS CQRNUTUS. -j. 




6. Value in microns 


10.15 


10.3 


10. 


45 


10. 


6 


' 10,8 


11. 





11. 


15 


Frequency 


1 


5 


4 




9 




14 


26 




27 




11.3 11.5 


11.7 


11.85 


12. 





12. 


2 


12.4 


12. 


55 


12. 


7 


46 65 


89 


55 


55 




22 




29 


14 




12 





12.85 13 13.2 
7 7 2 



Mi, 1 i ■ ' 


' '44-i-t 








m 










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1 1 








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ht-H 














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H- 




























H- 










































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4--| 




























































































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r 


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m 


























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1- 






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r- 




















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1 • 






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T 






























































































































































































































































































— - 




























































































































M- 






































































































1 : 


f— — 
























































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1 ; 














































: : 












1 
















































1 ' 
















1 






































h 






















































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1 






































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f- 




















































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1 


















































































































r 




























































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OF THE 

UNIVERSITY or ' 



IBEROSUS STRIATUS 




7. 



Value in microns 


13.7 


14.1 


14.4 


14.7 


15.1 


15. 


5 


15.8 


Frequency 




4 


9 


9 


10 


14 


33 




41 


16.1 


16.5 


16.8 


17.3 


17.5 


17.9 


18.2 


18, 


5 


18.9 


56 


42 


30 


54 


40 


36 


30 


23 




17 


19.3 


19.6 


19.9 


20.3 


20.6 












16 


14 


10 


4 


2 













OF THE 

UNIVERSITY OF ILLINOIS 




Value in microns 


13.0 


13.4 


Frequency 




3 


5 


15.5 


15.8 


16.1 


16.5 


53 


54 


51 


53 


18.5 


18.9 


19.2 


19.6 


11 


6 


2 


3 



13.7 


14.1 


14.4 


14.7 


15.1 


6 


7 


31 


30 


39 


16.8 


17.3 


17.5 


17.9 


18.2 


56 


39 


25 


18 


12 



19.9 

2 











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U. OF I. S. S. FORM 3 



I 

UNIVERSITY OF ILlIimOIS 



-26- 

IV. DISCUSSION 

While there has been written, in the past few years, a 
vast number of articles dealing with spermatogenesis and tracing 
"sex" chromoBomee as far as the spermatid stage of their existence, 
there have been few attempts to demonstrate the effect of this une- 
qual division, of the spermatogonial chromatin mass, upon the 
spermatozoa. Nearly a hundred articles a year are being written 
about spermatogenesis and yet only three papers have been published 
in which actual tests have been made of spermatozoan dimorphism. 
Yet the actual proof of the "x" chromosome hypothesis rests upon 
recognition of differences in spermatozoan lengths, separation 
of these lengths and artificial impregnation with the uniform 
derived product. 

Dr. Charles Zeleny amd Mr. E. Carrol Faust first recog- 
nized the possibility of measuring dimorphism of spermatozoa and 
introduced their problem by a preliminary paper on Anasa tristis 
in 1913. This was followed by a large list of determinations in 
1915. J. E. Wodsedalek (1913) measured dimorphism in the sperma- 
tozoa of the pig. 

There is practically uniform agreement that there is 
dimorphism of spermatozoa, yielding female and male producing indi- 
viduals. The ones with the larger chromatin content are commonly 
acknowledged to be the female producers. Inasmuch as the members 
of each group of this double spermatozoan population exhibit a 
wide range of head lengths, the two groups must necessarily over- 
lap and significant results in fertilization can only be obtained 



-27- 



by taking individuals of the extreme sizes. 

The data in the papers of Zeleny, Faust and Wodsedalek 
shows an expected bimodality in curves plotted from epermatozoan 
head lengths, The data included in this paper agrees with that 
of the above authors. The total accumulation of data includes 
some twenty three different species and its very extent banishes 
the possibility of obtained dimorphisms being due to any of the 
recognized sources of error. 

Six of the forms dealt with in this paper give undenia- 
ble evidence of dimorphism of their spermatozoa. The seventh, 
Passalus cornutus, seems to be unimodal and the eighth, Helio- 
drilus caliginosa, gives a dif f icultly interpretable curve. 

Passalus cornutus might be expected to exhibit bimodality 
inasmuch as there are a great number of beetles having "sex" 
chromosomes and practically none which may be definitely classed 
as being without them. However, the plotted curve is unimodal 
in character. This may arise in several ways. There may be 
only one kind of spermatozoa. This seems to be improbable in 
light of various cytological studies on beetle spermatogenesis. 
There may be two kinds of spermatozoa formed, one of which 
degenerates soon after formation. Morgan found this condition 
to exist in the aphids and it has been reported in the spiders. 
There may be two kinds of spermatozoa, the modes of which are so 
close together that the combined curve appears unimodal. This is 
the most probable explanation. In this form the modes might 
conceivably be at 11,5 microns and 11.85 microns and be practically 



-28- 



equal in height. The number of individuals of 11.7 microns for 
each group might be 3/4 of the number at the mode. The combined 
number of individuals of 13.7 microns in length would be 1 l/S 
times the number at each true mode and a false mode would be 
created. 

Heliodrilus callginosa is an hermaphrodite. Few ac- 
counts of hermaphrodite spermatogenesis are to be found but such 
authors as have published data upon the subject seem to uphold 
the view that two kinds of spermatids are formed. Zarnik (1913) 
traced the spermatogenesis of a parthenogenetic mollusc, Creeeis 
arioula. He found that two kinds of spermatozoa were formed, one 
of which was either nonfunctional or else degenerated. Schleip 
(1913) believes that the spermatogenesis of parthenogenetic spe- 
cies is the same as that of two sexed species. He thinks that 
sex is confined to the sex organs and that the somatic cells are 
neutral in regards to sex. 

Boveri, Kreiger, and Gulick, have also investi- 
gated the spermatogenesis of parthengenetic forms. Their concen- 
sus of opinion seems to be that two kinds of spermatids are pro- 
duced. It will be seen that there are several ways in which 
spermatogenesis may develop in parthenogenetic individuals. Two 
kinds of spermatozoa may be formed one of which is either not 
functional or else degenerates. Two kinds of functional spermato- 
zoa may be generated. One kind of spermatozoa may be formed. The 
curve obtained is flat-topped. This indicates bimodality. The 
modes probably come at 15.8 and 16.8 giving a ratio of 1:1.06. 



-39- 



ACKNOWLEDGMENTS 
I can not thank Dr. Charles Zeleny too heartily, for it 
ie his kind advice and endless patience which has made it possi- 
ble for me to complete this thesis. I also am greatly indebted 
to Mr. Hart of the State Entomological Laboratory for his care- 
ful identification of the selected species of insects. 



-30- 



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-31- 
V. SUMMARY 

1. Cytological evidence accumulated by the leading 
zoologists of the day shows that animals produce two kinds of 
spermatids differing in chromatin content, this difference proba- 
bly determining the sex of the fertilized egg. 

2. This hypothesis must be tested by measurement of 
dimorphism of the spermatozoa, separation of the two kinds and 
artificial impregnation with the uniform product. 

3. Since the spermatids receive different amounts of 
chromatin, the spermatozoa must differ in size as they are almost 
entirely composed of chromatin. 

4. It is the object of this paper to demonstrate head 
length differences in spermatozoa from single testes of the chosen 
species. 

5. Large range of variation was found in each case and 
the size groups could only be made definite by a large number of 
measurements and a plot of their size distribution. 

6. Five hundred measurements was decided on as the mini- 
mum number to be used. 

7. The general result was a bimodal or two pointed 
curve indicating the presence of two populations. 

8. Wherever possible the ratio of these two modes was 
compared with a ratio calculated from published figures of chromo- 
somes of the species. 

9. The general conclusion is that there are two size 
groups in spermatozoa of a majority of animal species. This size 
difference is correlated with a difference in chromatin content. 



-32- 



10. Acoording to cytological evidence the larger sperraa- 
tozoa are female producing and the smaller are male producing, 

11. The accumulated mass of data tends to eliminate 
the possibility of the obtained results being due to any one of 
the listed sources of error. 



18. Leptocoris trivittatus. The observed ratio is 

1:1.09. 



13. 



Reduviolus ferns. The observed ratio if 1:1.05. 



14. 



Corizus lateralus. The observed ratio is 1:1.09, 



15. 



Euschistus variolarius. The calculated ratio is 



1:1.04. 



The observed ratio is 1:1.09. 



16. 



Cosraopepla carnifex. The observed ratio is 1:1.075. 



17. 



Passalus cornutus. No observed bimodality. 



18. 



Berosus striatus. The observed ratio is 1.00:1.07. 



19. 



Heliodrilus caliginosa. The observed ratio is 



1.00:1.06. 



-33- 



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-37- 



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-38- 



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-39- 

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