Vidandg

March lL, 195)

Interpretation of origin and benavior of the eonvtrolling systems,
I, Revhew of previcus discussion,

1. Purpose was to show that svstcms present that control genic action in
manner not quite like that of Ds - Ac or that of a, - Dt,(waie. a e much
alike),

2. These other systoms, heave basic pattern however that is similar --=
control of genic action; evidence suggests that segzregations of controlling
units occur, “elotic segregations obtained and linkabes noted, Results
of somatic secregations similar to those of meiotic serregations.

II, Controlling systems revealed by the b.f.b,. evcle, Yoneluded that they
do not arise denovo but are present in the normal nucleus -- function in
some procise way during the develonoment of the organism,

“~ a). Somatic segrerations occur, changing the constitutions of the sister
nuclei woth respect to the controlling unit,

b). These changes result in subsequent controls of the action of snecific
genes,

ce). For the genes to become active, some change must occur at the locus of
the gene,

d). Some mechanism must be present in the nucleus thet init@éiates these
changes in nuclear organi,ation and constitution,

e). Believe this mechanism associ-ted with the heterochronatic elements of
the chromosomes -- primary initiators of the enanges that subsecuently occur,

f). The pattern controlling gones, the "switch" genes, etc., may be
expressions of their action,

ITI, ‘he recent studies of others showing the mechanism outlined in this
series of discussions®

1. Work of Brink and students on the var, P locus in maire: ilave come
to same conclusions regarding the controls of the patterns of variercation
expressed, ‘ave found the transposing units. lave found the twin
sectors of the types described in this series and have investizated the
progeny derived from them, Confirms the story here presented,

2. Peterson, in maice, has found system controlling varieration that also
appenrs to rollow the main conditions here discussed == the controls of
genic action; transpositions of the controlling unit ecmparable, in many
respects to Ac but not Ac,

ui ‘ * Ts » . 1 7
3. “he studies of the "Si" mutation factor in Drosophila by Ives and
Hinton -- suggests controlled chances at certain gene loci and controlled
types of chromosomal alterations that occur at specific loci,

h, Somatic crossing-ov +r as observed by Stern, Some cascs stronrly
suggest a controlled system in operation here to five cis
exchanres cCetveen certain chromosone contponents.

5. Many suggestive casos in literature. ‘the cythlogical cases are

most instructhve in this respect, Heview will be piven of representative cases
!
Ro
!

IV, According to interpretation given == that the effects described re-resont
those normally occurring in develonment -- we must account for their
appearance in the stocks used here,

1, If the vrocesses described of ecntrol of genic action occur in
standard streins, their presence not readily revealed, this is because they
are activé at particular times, controlled in tris respect, and therefo e
not observed as snecific gnits unless they ave called modifiers.

e. If the controls not regulated, a chaoite situation wuld occur,
regarding gene acticn, the resulation of develonoment, the maintainence of
snecies, etc,

3. he vattern controlling genes -= look very much like the svstems outlined
a), The patterns in flowers: strikingly controlled in many cas
b). The patterns of pigmented areas in the Lady Seetle -- again, “3
striking case of eccntrol of when the gene for pisment formation, or involved
in it, is effective,
oC) aarti ByBasde om
l, If cneof these svsmtems of control becomes "out-or phase” -- stearks
working at a different time in develonment, the results would be an»narent,
Altered times of rene acticn would be noted, Varieration would result,

o
oO

Qs
ot
o
D

5. *he mechanism of chance, AS
make it cloar thet once suen a ‘system be
continue to be so, and would r- i

genic action, elsewhere, whies. t>

utlined in the ctecyesiins, would
comes "out-of-phase", it would

ne aoner ‘prance of new Cuanges in
iid .e put ‘out-of-phase?

 

V. the interpretation cososi bed wovlc peaquire that in the normal develonnment
of an opcanism, somatic sesrorzations should cecur, (The nuclei should not
all be alike, @Arain, changes at particular gene loci should occur a

1 -

specific tyses, leading to nuclei with different vnotentialities for

action . 2
° GU Afi TEM on Mane gate

  

1. that do we know anovt,Gmnatie « segregati: ns? ~ About diffe: ence
genic action in different nuclei of an organism? ts therc any evi
to supnort the above intem meetations?

_ &« thoro is a vast wealth of 4
these arc ma°niy in the arce of inve
of it is purely deserintive, Hany o
xamined by geneticists because of +
not seen at all by biologists in ot
afe of vest significance, ine fie
opportunity is presse ime

on in sucnort of the idens riven,

cn imown as “evtology' and wuch
oaners in this Pteld are not
uichly teecrnical neture, Most a-e
dse The observations, “owev r,
riv been tS and a wide
nvestications,

3. A brief review w

ii) be given of the nature of this evidence, with
examples selected for i

Lliustreations, Will cever the foliowing topics:

Vifferences in nuclei of organism wit» respect to sets pn
Differcnces in nuclei af organisms with resvect to parti

“es prerent

DEP Le; ences with respect to component varts of the ehrorioec~e itself:

L “(a) Large sesments to (b) the rcuinutest seements known -= the bands
in the salivary gland type chr moscnme,

(1
(2
chronos
(3

~~ Oo ne
VI, Differcnees in nuclei of an organism with regard to the number of sets
of chrovcseres present:

1. 4ygote -- starts out with two sets of chronoscmes, one from male and
one from female, During developuent, chances occur in the number of sets
of chro-osomes within the nuclei, These are controlled changes: certein
trpes of cells having certain mecifie types of changes in sets, The
set may increase -- poly ploid nuclei; or the set may be reduced -- to neploid
nucle. fhese changes occur at specific tines during the develooment of the

organism, Also, different mentods of chance in set number orrur,

2 + nm { ‘ ce s ’ os
The increas:s in numbers of sets: By means of endomitosis -= no mitotic <
SO .
division ficures, Various methods of accomplis: ing this endomitosis.’ Fusions
a). the degrees of inerease: From tetra .phoid to 2,000 ploid,

b). The tye of appesrance i chromosomes Polytene type - salivaries
or diffuse tyvpe -- frequents; Or, combination of both.

The decreases in numbsr of sets: Some interesting examples:
fa pean 940 Y

a). Tail of Rana (frog), Epidermis is diploia (4° \tt* 4%
gpl yass, S8ee HIME is haploid

b). Sea Urchin: aA ! ‘reduction divisicn" in certain cells early in
cleavage Stage: 16 cell stage -- 1 micromeres vresent, These are 2n,

In division of ME CEOMCTOSs reduction of chrovoscone number occurs to hanloid,.
These haploid cells become skeletal components of oranien,

b). Increase of sets followed by decreasc of sets Culex, Very well
examined case to show the mechanism involved, Aoducti om oeet s during
metamorphosis poriod,.

VII, Differences in nuclei with regard to certain chror joscmes or the
complement: Loss of chrososonmes at specific divisions: RAMAEKRXRARERET
Non-disjumetions of certain chrowosomes in certain divisio ons; Cthor metoods
of acccormmlisuing the same,

1, Best workedcase that of Sciara: Outline on board,
Loss of chro.iosomes at mitosis
Non-dis junction of particular chremosome at certein division
Pasage of chrovoscme through nuclear membrane -- hignly ecntrolled,
the experiments that show which part of chromosome is resnonsible
for the vhenomena,
When is this effect impressed on centromeres}?
What mecnanism makes the impression so precise?

2, The B-tyve chromosomes in nlents, Very highly controlled non-disjunctio:
of these chromosomes, Occur at Syecific times and with s-ecifie consequences.
a). Example of this benevior -~ the B enromosome in meizes
iongram on bosrd,
-
Yontrol exerted at centromere vo cgion | © * cn
on st

f
X chromosomes in Sciara, The translocati udi

because - ‘eae pow of amu, Ho Lee ARE AMES
fee
“jo

n behavior of sets of chromosomes within same nucleus:
th. Wo, Coudtesad + 2%.
White . Monarthropalpus, Spermatogenesis:

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—7

IX, Differences in anvesrance of different chromesomes of a set in different
stages,
a). The smmll chr: oscme in salivary glands of White,
tN " i it "l ut ite] land

b). The alternation in condensation of particular chro-osomes?
a
Tye TN 2

X, Differences exoressed by individual parts of chromosomes in different
celis of an orsanisn:

x

A). The chronocenter regions -- heterochromatin in Drosophila -- too
well known to commnat on,

2). Condensed versis diffuse condition of segment of a chromosone’

a
TMG Tuye v
wb "od
3). Most exciting recent case -- that of Pavan on Rhyndhisciara -- two
different species studéed,

a), All eggs laid at one time -- within lS mins.

b). “early all lerwe of one sex

c).All develop at same rate - 2 days to pupation. Common puparius formed

d). The changes occurring to snecific bands at ce tain times in ,
development in the diffcrent tissues; change at particular band svecific
for the tissue involved,

e). One tyne: at certain time, increase in D.N.A. and swelling of
particular band; Separation of commonents of the polytene bands --
arge Puff region formed, At certain stage, shrinkare of puff rerion,
Return to dense band, but now bad very much wider than before the process
started, Increase in D.N.A. remains ¢ Interpretation: Multipliceticn of
genes of tris band, specifically. Yifferential beh-vior,.
XI, Conclusions on nuclear compositions in organisms: Wide differences
among the cells of an orsanism with regard to the nuclesr cormcsit. on,
Differs with resnect to sets, parts of set, p:rts of chromoscmes,evenparts of
so small as a band in the salivary tyne Chrenmo somes

“echanisms : Lifferential behavior at certain mitoses: Losses of
certain chromosomes; Non+disjunctions of controlled tyre at certain
divisions. Passase of chromesome throurh nuclear membrane ¢

ATI, Differential nuclei resulting from specific divis'ons: on genic
level,

1, “aramoecium -- mating tyne reaction:

- '
c t fe ‘ . . , ta WM Ore ely aad bor. VAA
re. , 4 LiMALCA \ a
(") () ate -] pegcggbur Arb Lag. Uderesaiti ts ererirg cel -

me ee

2. Tetrahymena -- mating type reaction: Same as paramoecium except the
division is not as precise -- first or second division of macronutleus ,

s

3+ Case in Glomerella where mating tyne "mutation" occurs at specific
time in develovment of ascogenous hyphae,

XIII, The effects of svecies crosses: Often quite bizaar,
1. Changes in the chromosone cormlements Various polyploid conditions

reductions, occur in tissues that do not show sucn reactions in the
individual svecies involved,

3

2. Case of Metz in Seiara: Cross of S, x Ss,

the mosaic salivary gland?

3e Species cross in Nicotiana, A. x B, Backcrossed to A
A.A B comlement: B chromoscmes eliminated somitically.

3. Altored cenic action often noted; Variegation anpears,

Ch oA LE

XIV. The constitution of differentiated nuclei -- as shown when these
cells forced to divide,

a). Treatment of root -~ chemical (indol-acetic act@d. ces not affect
behsvior of chromosomes in the nuclei that regularly divide. Forces
nuclei that are fully difforontiated to divide. Theh show a variety of
chro osomal aberrations, One type observed ine ell that had undergone
endomitotic divisicn:
~6-
“seh Shi.du wal ok- Cry ( au

AY
// \ H jj \h Sowhereane 106 Sot

The effects observed are very much like those produced by Ds,

XV, Ds type behavior in other organisms than maize?

1. “ete - Sciara, Certain chro “osome, certain band region -- break
seen in all salivary gland cells. “howed that tois was penetically
controlled,

2. Hyacinthus - Darlington, Break in a particular chre mosome late in
development of the snorocytes, Genetic control

Be Vicia - Levan, With certesin treatment, break occurs at the
nucleolus organizer quite predictably.

l.. Dreyfus -~ break in arm of particular chromosome at certain division
in the svermatogenesis of e

5. Gatcheside - Venot-era, His case of position effect following
translocation: avidence much like that found for Ds Chrorcsone br aks
chromosome duplications and deficiencies of certsin rosion of a chromesene
in the transloention com lex,

XVI, Conclusions:

1.
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