Pl!RMA.'.OOGl!S»llS AND OOG!BmIS OF BUM>HCRUS CON'.OOR'l'tJS,

N 1 '!ODE PARAaI TIO IN THE WlUSUM OP SBXEP

A:pproved:

AND OTBIR RmaNANTS

M7"Hl I'. itenderaon

Majo7 Thesis Submitted to the· Paoulty ot

The Virginia Polyteohn1c In.atitut• D•pa~~men.t ot Biology

l'or the Degree or ster ot Soience

in . Biology

Dean

stipervieor

.Tune 1939

ead otnepartmen t ot Biology

ll!lTRODUOTlON ••••••••••••••••••••••••••••••••• * .............. . l

RltVIEW O ••••••••••••••••••••••••••••••••••••••••

li'tTI..S'.t'l G TlON •••••••••••••••••••••••••••••••••••••••••••••••

Procedure •••••••••••••••••••••••••••••4•••••••••••••••• 5

i1eth0d• Qf Obtaining peoiin•n• •••••••••••••••••••• 5

Hiatologic 1 Teobnique •••••••••••••••••••••••••••• 6

RerJ.lta ··································~·············

SperJ11.togeneeia .............. , • ,. ••• ., ••••••••••••• .• • 9

eo.geaeaia ••••••• ~ ••.•••••.•••••••• ,. • • • • • • . • • • • • • • • • 11

••••••••••••••••••••••••••••••••••••••••••••••••••

lD CONCllJSl.ONS •••••••••••••••••••••••••••••••••••••

DliS CRl P'l'l OJi 01' PI •. TES •••••••••••••••••••••••••••••••••••••••

••••••••••••••••••••••••••••••••••••••••••••

--••••••••••••••••••••••••••••••••••••••••••••••••

13

15

17

20

21

' INrHOWOl'ION

Baanonchue contortua. otherwise Jen.own 0.8 the "Barber Pole

"The lre o " or imply aa the l rse t ach oriii ot heep,

h • been he subject ot ch ac1ent1t1c tntereet tor the p at

que:rter or a century. Consider ble work baa been do~ whic hn es-

tat>lilthed tbia puaeite o.a e. te.cto:r ot econ.o ic tmporte.n.ca end its

presence aa pathological agent ln th a sum. ot inanta h

generally well :reoo tzed. Th1s 1'01'11. 1e one of the l rg et ot the

Strongloid ro1,1 , me suring 10 eUmea u illch 11' the t ea nd

•ell Jm.oa. Tht 11'e:ra~ure doea not, owe'fel', rt••al u.J .ork which

hae been done 1n oonn.ect1on with the deYelopment. ot the reproduot.ha

cell~ of this epecie •

ln the reaent work a atudJ h•• been made ot ape togenee1s

and oogen aia wUh JH atten-Cio d1l•eote4 to 11etud1 ot copulation,

tha »aasage of the ape cells ~pon entrance into th vulva. ad the

ane.tomtcel loc tion in the f'esnale where the sperms are collected

tor the purpose ot tertilize.tion ot the OY • o 1etologlc section,

4ep1oting the ntrance ot the ape · into the ov • h • ee obaerved

but developmental etagea immediately following \hie penetration ot

the ov• are ebofm in thin poper.

Even though considerable amount or me.teri l b been

examined which has afforded opportunity for tudying the entire

2.

reprod.uoUYe organs ot bo'h male and t•al• worms. aeYeral develop-

mental etagee which hRYe bMll rePorted •• ocOUJT1ag ill N toda,

have not b&•• ob&eZ'Yed ln this stud1. Sutt1cient eY14ence, however,

1a herein preaented. whieh illdtcatea ••rf clearl7 and det1n1tely the

o?mal ohram.cu1<•• number ..

A oompll tlon or pu'bl1catto . deau. . 1th he ohromoaome

number or paraa1ttc aema'Od•• ie &Wl1la le 1 ileon'• (10)

Tb• ,2!ll. .!!. De1elopn.t !1.4. §en4tt:r:, 're» espedUe a comparla:on

ot :re•ulte ot other .oJ'kel'e a.nd \hoee de•orlbed 1n th1a pa.,_r i

ll•t ta recorded below:

Author's N-. Dete -'en Benedin l8S~

'f'9l'1 l88V

:.ont1tomeey 1904

Edwrds 1910

Gulick 1911

Bo'fer1 ed Schliep ltll

llulsow 1911

'.ejdoTalQ' 1912

Goodrich l9lf

al ton 1.916

(4) 1gs:

Spee1oe

cana blnlens

Aecar1s . msalocepl!!J..,e. uniftlena

Parago-rditla Yariue

ucaria 1umbriec>1dee

Hetera.k1s Yeaioular1e

Aeg1oatomum (Rhabditl•) n1groTenosum

Anqraoantaue 9at1

Corcltua \olmmus

Aacar1a in.CN.n'a

Aacaria cania caris tel1s

AaCtl?-18 sUllla

caris vituloru:m

*This does not a.ppear in Uilson (10}

Chromoaome $umber

,;;; 2 , ZN _. '

• 8 in tae mela

=?. 2N=-.l4

2N=13, !i =-19 and 24: 1n tho BIAl&

=-9 1A \he male. ~10 hl the female

==n 111 the male ( Schlle ta tb.e .U ff; 5 and 6) -

=12 1a the teal•

=-II ill the male ::12 ill the t91Ul e

:::::.'., 2l'if""•

Jl'.=14'. and=2l 2!J= 35 and

2N:::43, !~19 and 24 in the male, .R"'24., 2N=•e in the temal

2lJ.,,.9 in t!le male

p •

Ct ur•

the cour e of t ta 1rt'eaU t10A ee'fe l heaY1~ 1n-

tested. llh•P were el tered Jl4 the abomaaa were tnO'fed. • oon-

\enta or • bcDaaum •r• .a e4 out into jar ot pb7atolo 1eal

• lt olut 1o1a at 4'00 o. The stem.ab. ea tbe split longUudtnell7

wh1oh ha prntoull been filled wlth phJe1olog1cal lt 90lut.1011

t• e. ebDrt time the woma tree the.eelfta trom the

muaoaa end deec ed o the neck ot the tunnel ere they were

• 'rheM woisu were then picked ut

•coordiog 'o apeo1e• wi t)l the aS.4 ot a biaooul r ado co~.

tixed 1naed1otely, t1tt7 ot the•• were

re t n tl'O.ll \he Bae n ap ratua or t'l'O

he auooea and placed ln Ringer's solution end kept ta atu- bath I

at 400 c. trom three oriu ot eacb aex ere

t 7:00 P• m. a tue at 1ntenals ot one hour ea tor p_.iod

ot t .. l•• houra. T 1• • done in Yiew ot the oaeibillty that

1toa1a and metoala in th se wo1'Jna might occur 1D. cycl • or at

6.

Hlsto,lgsloal. Tacb!U9

JTevlous unpu l iahecl work 4oae by ol en ( 6) 111 thie labora ..

to17 ladioat 4 thll' hot 'aboll1 ?00' C.) Cano1'• f'1xe.t1ve a Ued

tcr thie particular type ot tie e, 1 t t tbe "° tued nearly

strai h end othentae l as co ort . • e tin ct1on na .

-1.lowed to oon'Ua.ue o'f•r a piertod ot tt'el'f'e boura, t en remov nd

atoncl 111 '10 percent •thJ'l alcohol..

'Gpon. l'8Dl0Val tro ?O p1rceat alcohol th w s were plaoed

111 4iaphanol tor t"rom to\I!' to eb hour•, 1 or411r to ortea the

·Uo e • o •nvelOpiJe each rWa&l • Zirkle' a ( lO) t hod tor the

prepa:r ~1ou or •0041 plant t1aaue,. bJ' tu u e ot •bUt7l. alcohol

'dur-

iq 4ellydra'Uon e.114 11'1.t1ltntS.on. In thle aa.e 'the worme •r•

atarted 1D the 60 pe:rcftnt aoltt ion ond. kept onl7 .:50 1n'Q.tes 1 each

ooncentretion.

From the a baolute b\ttf l alcohol tbe a~oimeu er ftll!?ve4

to a mixture of halt e aolute but7l end halt paratttn, (:S0-520) and

allond to rematn tor h'0:1D t to our llO'l.ll.'8.

to sort (f,0-520) pare.tttn, tor rrmn t\'fO to tour hour , the into

tin 1n 1ob. they we 1mbe e • Hard (56-58°)

tmre.ftU. was used 'but wa.e found to be too br1t~le .tor thi& type ot

tissue. Ti au8118t e.l waa esn loy d a.a an tmbed 1 d11.ltD but

'· d1tt1cultf ezpertenoe4 1 maintal iag the 4••1l'rlble oonaia\ Cf

tor seot1on1ng.

eara were made tram o•rated gonad• • rdi~ to Btllli •a

lro•-e.eeto-camtae •'bod. 'l'hiB ••tb.04, aecordl to t • ~· 1que

emplo7t4 lll thie atudr, •it tlOt aucc .. etul.

ale and t ale apeo1mtH1• were sectioned t ab: S.cron •

11• tab thtckn.e111 was autt.abl• to~ 'the ale ge celle, th• tamale

erm cell• ••re rendered. too thin tor proper atud.J. ,.._le •o a

cut a' te~ and irtaen 1orozaa pro•ed 86tietactol"1 tor ex 1nation.

Cr<> nctiollS end lo11.s1~ud1n l ee Uons were de ot both sexoa.

The 1ong1t~ inal. eectiona ot the male• were ecidedly au.perlor to

t e orosa aeotioaa due to t :l:le rpholOQ ot the teat•• nd their

loce.tton •1th respect to tbe al.im ata1" oan l. 'fbeee orgt.U1s parallel

the canal anteJ:O•poateJ"iorlJ thua attording complete eeotio.ae in

which developme \ trom the •erlieat e cell8 to the ~ure a erm

uld be studied. In ihe t ale the oi"Oea aectiona e ed beat

aulted tor study due to tho toot that the O'farii.• ot tb.e t ale form

a pire.l aroUQll the alimentary ca.1. henc eectio cu\ lengthwiae

attordad e. lees number of ••11• 1a one plen.e a.ad the materiel was

more easily torn in preparation.

The sections were mounted o elide in a rtal order and

allowed to dry tor twenty-tour hours or lo er betore they were

e.

atained. They were \hen nm through two cban,, es ot sylene tor ti ve

ainuies e cb. 1 \hen t rougll the regreoeiYe ethyl alcohol series tor

one iaute e ch to d1at1lled •ater. 7 the w ter they were

l"du.te 1 2 .5 percent iron al tor thirty 1Autea. she4 in

tap water tor tit• .. n mlautea d ~snoYed \O B1denha1n•e h a\oxylin,

to which one drop ot t rgitol ff ha4 beea dded tor eY 'l!1 a1xty cc.

or ate.in~ tluliJ. Th tul'gitol • us ttel' ce'• (5) method.

•1th axaept1on that thirty 1aute• time as ell.owed tor iat.ni •

e elidee were Ulen rinsed 1a t p \er tor tit~••n in t a,

4 stained 111 eaturat~ picrl.o e.ot eolutl.on. n.til dUterent1e:tion

ttainecl. The ection w re t •A wa.ehed 1a l"l.UU'li , ••ter r

r1tteen in tea, run ck up t e· alcohol aer1•• allowing one inute

1 eaoh couoentl'8Uon wit except1oa ot abaolu'e aloohol in wnloh

they were kept for fifteen minut , t Ch .e et le e tor tive

inutea ao n:re al.load tor clearing tbe •c\1ona fro when e

theJ were r "fed &.nd mounted in b le thod ot

d b.J'Q.ratton u deecrib by Guyer (4) u ue d on aeYe l slid.es,

and while th• method a Yee tim 1\ apparently act.a. ao &wh. t

bleachin agent, 1nce slides so treat either pre •nted lees

dU'teren l.atlon or the at-1.ill appeared: T•J7 •eek. A tot l of thirty

le l!Ol'Jlls end titt en t ale wo · s were examin•d 1 the study

h rein described.

9.

Re1ulte

SJ?•ma~os9ea11

The 4evelopnent ot the pr1m.ai7 epeniatoqtea troa 'be

apenaatogoale baa not, ln thla e'ud'• b .. n. ob en "1' u.y degree

ot 4$CU1'ac7. It ia bell••• that the \nna1toJ'7 atagea 11l'fo1•

Sri the development to tlllJ. sperm in §!emonc=h contortue en

exceedingly brlet en4 that the reat1ng etagee •hi will be 4eaor1bed

:re th• r at1 ategea 1cli wtll • tecribed e the reati

a.tagea of th• prlUrf apeJ!l'U.tooytea.

froat. twelve all ea o't loag1tud1nal aeoUone ot le p c1 ens

tn which each MOUoa lhond the eats.re le114th ot t e tutta an •

tberotore tbe 0011Pl•t• deve10.-11t ot t.be reproduoti•• tile. lt

1• apparent that 'h• d·&'Hlopme:nt proceed• trom th• e.nterlor portion

to the po tel'iOZ" region Of the tea·Ua • The Oell• e.ppear ~ery all

ia the terior en of the teei1a. ft• nuoleu 1 - de p ot •

•ll&l• auoleolus and 11111a tbl"ea4a. Theee etl'Uctur • H located

juat •1th1J>. the nucle«r Du1111brane aad atain lightly a co pared with

the later ateges ot their de•elopaent. a pl"Ogreasl• d Yolo nt

1ncreaoes the cells bec0tne le.rger, !QOre elongeted, po seas a

definite nucle r eml>rone but as 1•t no cytopla io numibr ne cv11 b

observed, 'Figtt. l, 20 nd 21. from this 901nt there follows a

gnd.ual e'.bange ithout enf indication that a mitot1o d1Y1s1on tokes

place. A eyn.cy·Uum pers1.ete, the nuclei become ore apherioal and

10.

l•rgel" EJfld the llnin threads hee.'fter cl tb rather lerge gl'Bll.Ulea

arranged ittegul rl7 Oll these threaisa. other workers he.Te des•

acr1be4 theae granule .. laaTiDg art en t b l r • nucleolus.

This aeama hiehl.7 pti0ba le elnce with the ppearance or these

g"ftJIUloa the nuclo lus 41aappeare • fig a• 2 t 3 nd 2 • Condene t io

ot chrotne.tin terl l, 1g. a. proceede, tormln.g he&"f1 ep1:r e-lS.ke

st1"Uotur a 1.1hiob a.re atmtlar 1u app~uoe to structures ct.ch ba"fe

' bee eaor1be4 1n t e lUera.ture e the 41ploteae te.ge. J'rom this

atage there deTelopa lame 1ately tour l rge dia4 4 one a~ceesory

chJ'romOeonie, J'iga .• 4, 6, 6, 'I, and 13. The n.a'ture ot t e e d.iada

btJoome re apparent u tih"7 ttesol'f• into eight ll monads end

oae Ol'l!t body Which ob'f1owsl7 1 the 1ex Oh so e. 11 • , 1a a

Calle~ luo1da Jl'BWiDB dep1ct1 tbeae monade. phot 1crograph

oould be obt ined tor t i& deYelopmental •tage at a

ditl not appear lD oa• plane. J'rom .Fig. '• on to the complete

<le'felopment ot the sperm, no aex olltom.Oeome can be itterent1 t.ed.

l1'1g. 9 • ahowe a di T1,dcn er a11aphaae or etoaia. tages, were

ol:.laened frequently at1ler the gJ'IOupe ot nine s :.e de their

appearence, t 'hia obaertation was not entirely con iatent as

there appoare4 in some aeetiorus, group ot tour and ttve chromo omea

witbout 8Jl1 oYidence ot tbs presence ot these d1v1a1on eteg,es, . .

:Figs. lO and iu, These gzioupe, because ot their co t1nuous occurcnce,

1n the m tertal x 11lined. are regarded as reprcoenting the normal

11.

b.aploid cb?'OUIOaome nuraber. J'ig_t. 12 • 13. 25 and 2& • repr ent. tbe

Cbaracterbttc oad•nse:Uon ot theae groupa ot tour 81'14 t1 Te cb.romo•

eoaiea into ape t14e •h:Lch de'f lop int~ permatoztni. :Fig. l• and

7. In two al.id• tl'Ola two tttere-nt •o • th•re -.ere toun ebro ~

soaea aberratio~•. laYol'fing a dev1at1o~ from the r$gular haploid

number ot tour 4 ff.Ye to • r•tner un.blllccea condit1oa of three

and six. This b 1agr ettc Uy 1lluetre.ted 1 Fi • 11. w ile

Fig. 2 , 1 a pJ:lO tcrog:t ph ot tbia bno l ooa.d1t.1on. Thi

etage waa observed ui te otten ln t e reduct ion stage but theee oei

groupa or aimilar onea CanJlOt be tound tn that stage •h•r• tho

normal gl"OU.PG are coad•n•ing to torm the spe:rmatld • ln. vie• of

th1a it ls bel1eT d tnei those abnormal gl'Ou s ar• iiher absorbed

or do not mature io e ome tunc~ion 1 aperrn.

ooenes1.,

In -th1a atudy the in•••t1ge.t1on relaUng to development ot

ti. remale gem cell ha• not been oom,pleted, bu• :the eta.gee aho••

1ng several 1ntfll'est1ng ~ha8ee ot th1Q developttent are desr.ribed.

'the very 8.<irliut cell3 ~re ch. r cterized. by pe ring

~longated and per,endlaular to tb long ~xia of tbe wo:im;, ~1th -

lief'inite cell wsll, and a oo:mparat1vely am!lll heavily at a.ned

nttQleus. Tile mornholQg<J of these oell:!t was obeerved. to change but

llttle with the exoeption th.it they dp~ear mueh larger as the

/

12.

ovary approaohea the aem1nal. v .. 1cle. D1tferentiatlon ot the

chromatin mter1a1 waa obeened in the pr11•J11U"T oooyte. np. 15

&nd 29, eho• tbli arrangement or the dyads 1th1 the nuelear

membrane.

1th the ~ne ration or the egg by tbe apena reduction

div1aion or tbe tol"!ln ion or polar bodie• 1e uaually et11t1U11lted.

In igs. 16 and 17 reepecti \f&ly there r tteen two bodiea •hi oh

a.re datini ely co po~cd of' !*iv cliro ·~ o ,oe. • nd d«t pUe the f ot

that no epei•• ' ppear in the o ll.. or 11u:leea ftnl1" oh1"()mat1o

mterie.l which ' gh1. rea$1blt!.< pronuel • o:!' a •P<trtri, there ia

no other altern tive to follow but to conclude t t ineso ~odies

are ~olar hodi••• Th9 tive e roinoeo a in the lower oun ry, 1n

Fig. 17, indio t~ that reduction d1v1e1o ha.e btten ao ompliahed.

Tile celle in Fi •. 18 ed lt were looat~d. in the uteJ:"U•

•lightly d1atant drom \he ~· in l receptacle. The position or ~h••• oeli. clearly indicated that rert111~t1on hsa been acoom-

pliehed. 1'1~. 19 1.e here df; cribed t.ts depicting the early anap ie.se

tic divis'i. on ot' ,he f4't:rti11.zna egg. Twenty

ohro:mo8om.a a.ppeii:r. e.:i.n:h d.au~htM'" oell reoe1v1n.~ ten. lJ'I . .Fig, 18,

i• seen )Olar Vl 0 ot ~ · si r t,o t t in 11r,. 1 • Fi •

30 od 51 are m.lc.rophotograph• of the ea.me oella deacrHled in t'igs.

18 and 1'9.

l~.

S2 Plate V

COlVIATIOJI

The pbotoa1 l'Ograph, ig. 32, ahowa a ale an4 t e

Theae sp cimena were obtained t an abom.aaum

ot a sheep 1n. 1935. Th re tixed in hot '70 percent ethfl

alcohol and untad 1 gl7cerine and later photographed by Elizabeth

Conn.er.

The phenomenon 1 tairl1 o n where the a sum is

heavilr pe.raa1tize4. Fixation otten osu ea the wor to eper te.

During the eaent inTe t1gat1on 1m1le.r apeciJnena to those des•

cribed abova were uooee tully tlx d but 1pulat1on or th worms

14.

in PNp&t"O.t i()n to:r: imbed<U.ng pt"OVlil <.Uuu tr0"..i1.1.

!be .. • 3 , ill t t-.s dt>ti 1tal7 the u o of the

b'lU"at\l ra1s snd th pio\tles 01' h wl urins co.vu t 1oa. •

l&ttar re 1zwerteu ll i :to hl'J VEigi hilo tbe bux la puebe

baek:~ rd and clo elf appl1e4 to tho body of the t ~ ~ot1olla

th:ro h the ,1>011tor10. riegian of th.i mt:al sho1.1 ·n t th picul •

... a hollo\T.' and lso th t, !.'Yh~ t 1 · r pl e.a. t th;,1 , uJ 1e

ahown in the pictu.cc thnt :t0th r holl ~tube 1 thua :ormed. lt

could not e d bol't:l,in t::-am u tudy or th · tt¥oC!m~n a to t &

method by wh.icb eperme r'1 t!'tluterrcd rro the le tc th t ale.

Great numbero ot opetm appear 1 eoction.• cut thrcU«h the

ovij c'or• a d ey 'bo found ill the anti~ latD th ot ho uteru •

:'hey are partioul rl:r plentU'ul. I.ii tbe o in l •eldcle and . here

'"1le oigan ia co -tr1ctt4 it S most p:ro'b bla th t hore t one

celled ova are tert111~ed. f«o.e:le eelle •h1Ch haYe pas ed th$

aem1nal. •eliole an4 h :n entered. the uteru.e b.e."fe alr ad1 b ~ to

Mgm.en1t, hence 14'; :Le quite &'fident tbet t.b.18 nioulor rt Of th8

fee.le re roduot1ve system 11 the ite or tertil1281iion. Since

th1s structUl"e is ver:v abort U ta exoee41ngly ditf 1 t to o'bserv&

tbia proceea.

i,!rm&togen~i•

l . 1a.t1oA ot lon aeot1ona, • ,gitel aeot1ona, n.4

<1N•• •ectiona or t e le ematod•• Hae<>Qhue cantortu• indicate

t ' the develoiment at t- au cell• proceed• t'rom \he anterior

end: ot the teatis oaar4e the poa\er1or end.

2. The tirst tne or rm c lla w iob could be cletinitely

recognize4 ill t ter1al at died ue apermtoortea ot the tint

or4eJ' .

3. ?10 aox ohrot'Oeotaa oould be reooeair..ad in the reating

eta.ge. of the p~1t.iary aperms-too1te.

• T lepto ene ia ropre ted by nwnoroue tine,

we ·1y s 1 1n 11nin thr•ads a ehowu in Fl • ! .

5. '!be d1plotene 1t preaent i• so tranaitory t t 1ta

nature could not be determined •1th an7 degJ!'ee ot aocu1"tloy.

6. The ~ohytene et ta repra anted bf ns. 3. eae

h T'J' threads do not apl1 t but reac>lY• t Nelvea quickly 1nto

hnTY and tbic er obromoaome bodi a. ro theae bocUet tour dya4e

re tormed and one acoeeaory o~ an•cbrc osome designated as n

X type ohromoaome, Fig. 'I .

7. From the dyad tom• n ne mona.da l"fl8.\ll t whtoh separ te

into two groupa ot t111os ana foura. These oups ot cbromoao: e

1e. condense to torm the ape1'2l'l9.tida.

a. QbrotM>some aberration& ere obaerved to occur in

which groul)a or eix ttd thr•e ch?O • were aegre - te4 . 'thee•

groupa ore not observed to ~orm ape . t1. a .

9. 11 tle ~oPQlatlon a tor many yea.re !>Mu kn.own to

occur in thi-1 sp e1 ... obaen ion ot thia pheno11eAoll ha• hot

be n reported. Th• aboY• ob•el"V tion h a been mde in th ••

laborator1e• 1 l93S 1a 4 ring thi• inYeat1 tion in 1938.

Y&gina or the t le 1• detini el7 entered by the 1p1oule1 ot

the mi..

10. There it ~o e•tde~ae that thal'e ia ~ diYiaio~ stage

in the eeooAd!ll'Y epermatocytee to toi-m the 11permtid1. O eena·

Uona, howner , do ind1o t• ' t eperms.tide r&ault by condnaation

ot the ahl'omoaomea 1n the aeoondary ape tooyte• .

Oogeneeil

u . ThJ;"•• stages in the oogeneaiB ot ti. t le ematode,

emoncb.ue oontortua were obsened, amel7 the primary ooayte

depictin fiTe dyad c1romosomes; a secondary oooyte oon 1n1n

one polar body ot tivo ohron-oso ea; ~n e i-ly an phase e or

aoma.tic d1v1aion juat ~fter fert111r.ation. 'l'wenty ob.ronoaomes

appsar in this e'4 , G'lCh daughter oell receh·ing ten.

1?.

nESCiUPTl.Olt or Pl;~ ------ .

Pt 1'B I

Oder« Lucid& The 1t1cat1on ot e o lu¢1d araw1nga a.NI not

stated beo uae accuJ'S.te r1 re• oantiot be given aince the dra.winga were tranetered by rne llcd to the plate and the plate. photogxo1phe4.

Fieun 1. Oell ot •~l. n•t1ng at

oyte found tn the nteriot" en4 ot t teetia.

P1gure a. Celle ot late!" at4ge, toun.4 more ,POateriorlf

1a the teet1•, 8h&w1ft the 11l0l"Ph0logioal t\ovelopnental P1'0C1•••

o•er tho•• in 11gu1'9 l.

r1gun1 '· 5, e. poaa1ble o l'OJnOaon:e tor.at1ona wb1ch "9J'e observed ia the aeocnd ry

epematooytaa witb tour 4yada an.d o • sex•obrotrJDaome.

Fi re a. Form.a ot uine mo 48 •hie reault tro the

?iguH 9. 'Reduotton diviaton •tagoe obsette4, rewlting

trom tbe groups of nine monads.

Figure 10. Seoonau.ey ·~•tooytea formed through re-

duotton divi ion ehow1ng noJ.Tal haploid nu btt:ra.

Figure ll. Chromoaome abenationa resulting rrom a,bnonnal

reduct1on division.

18.

Pigure 1a. condei:iaa ti on or the normal haploid munber•

ot chr01D011ome1 1how1 four ~d ti ve in a-ao

Figure 13. rurther aondenaation ot the ohromoso •• w

toftll inaature ap•rm, tnowi~ 6Llao • deti~i te o•ll •all.

figure 14.. tun •pem. ..

Figure 15 . Prtmry 0001~• ahowing ti-re Clyada .

Figure 16. Seoond~;r oooyte ebo.tn 1>01-r body.

figure 17. eoond ry oocyte sho1t'1ng a polar body and t1••

tunot1oul ohJ:ro110ao•• ·

J'igu:re 18. Po r view, ot earl1 enapha•• atage o~ somatic

division Ju•t attet tertiliz tion.

ieure 19 . Long aec-tional vi .. ot early WUt.Pb ae at-.

ot aomatio d1Tieion abo•1ng '•~ty olU'omoeomee .

PUB 11

Fhotomicrograp (x 3500)

igarea 20 and el. rly reat1ng •tages of ma.le germ

cell• aa ta shown in Figur• l 4iM a. '11 gure 22. Later • tagea ot ger cells s a own ill

igu.re s. 1gure 2~. D ad

4, 5, <>, and 7.

upa which rs represented in igurea

l9o

PI.A.TE Ul

igure 24. Haploid groupa ot tour nd tiTe oh:r0Jn0eo111ea

• le •hown in Pi r• 10.

l'igure 215. Cond~?lB Uon ot the groups ot rour and ti v•

chroaoeomes, show in 1gure 12.

Figure ae. CondeJ)Jled llllillllture &pel"'!!'.8 •ith 4et1nite 4ell

wall.a, ehown in Figure 13.

Figure 27. tur• aperm.

PLATZ lV

Figure !'1gar• 26. Reaults ot bnc;urmal redu~tion d1via1on,

Ohl"OtlOaOX!le abetrationa, a.a sboWQ in Figut-e 11.

Fi re :e9. Pri ey oooyte showing t1v• dyad• a.a in

FiguJ:e 15.

:rtsure 30.

Figure 31.

oell trom whic 11gure 18 waa taka11.

cell from wbioh rtgure 19 waa taken.

PLATE V

Figure :S2. Described on page 13.

, _ 0 1

.. ~· ~- .. 10

.... -, .. ·=:· •••

s~·· .: 11 •••

.,·, .;• 1 •. '

·•' . •. . ·;

\2if_,~··'· ·,,., 15

18

t:·

ff)JJ~' , ~a·· .. ;t ~f_ :. ;: ~.. -. .. , ... ;' ·~--. 3 · ~~ ·-· · .

.

1:: ti'•. ... .~ . 8

•• •¥ 12

/ '.

16

19

PLATE I

~) c~ e

14 @

\

20 2I

22 23 PLATE II

..

24 25

• •

26 PLATE III

•

• •

• ~ " •

- • • •' • • • t;. ••

28 29

. ,

,

' . ' , \

•

30 PLATE IV

31

20.

robe authoio wtab.ee to expreea h1• i>:rotound appreo1e.t1on to

nocto:t- • L. 'fbJrelkold. tou bis tim , ottorts, un4 suggestion• ha

so willin':l y ge.ve ud W"1tbo it •h.1nb tbis wol."k would il:lOt ha-ve been

doiie.

JLcknowled oma.nt 1.a ad• to who ou:ppl1ed

t photogn\pl\1o p:t te ft>r figure 311 and tor typing this th.esia •

c nowledgement ta lso 4• to fQ't'

her valuable help and euss et1ona in xe.king the p o•omorograph•

in thie thoaie .

'l'be author wiah.ea to expna• his appreoi ti.on to Dootor

1. D. Wilson to~ h1& •noouragement and helpful a~itlotl!UQ.

21.

BIBLIOGRAPHY

ea.

'I. Homedea, : •

1933. E11tu io del aparlto ClU."O•mQ•KmllCo loa

( GOez ) • tlev • d lU. • n.,

a. Schliep. •

1939. Tb 1nc1plea ot Oelleti ea. cGr ..

10. ilson. • a. l 20. Tho Cell 1D V lopment

ll. Zirkle, 0.

ac1one ?l: 103-104.