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Kalthoff / Second Edition Contents 1

ANALYSISOFBIOLOGICALDEVELOPMENT, second edition

Klaus Kalthoff / University of Texas

DETAILEDTABLEOFO!TE!T"

#Basic Principles)

(Key Strategies)

$A%TE&' /A!AL("I"OFDE)ELO%*E!T

1.1 The Principle of Epigenesis

1.2 Deelop!ental Periods and Stages in the "ife Cycle

E!#ryonic Deelop!ent Begins $ith %ertili&ation and Ends

$ith the Co!pletion of 'istogenesis

Poste!#ryonic Deelop!ent Can Be Direct or ndirect

d*lthood Begins $ith the +nset of ,eprod*ction and Ends $ith Death

1.- Classical nalytical Strategies in Deelop!ental Biology

E!#ryonic Cells 'ae Predicta#le %ates in Deelop!ent

nalysis of Deelop!ent ,e*ires a Strategy of Controlled nterferencesolation ,e!oal and Transplantation of E!#ryonic Parts re Key

Strategies of E!#ryologists

1.0 enetic nalysis of Deelop!ent

1. ,ed*ctionist and Synthetic nalyses of Deelop!ent

$A%TE&+ / T$E&OLEOFELL"I!DE)ELO%*E!T

2.1 The Principle of Cell*lar Contin*ity

2.2 The Cell and ts +rganelles

2.- Cell Shape and the Cytos3eleton

Cells Change Their E4ternal Shape s 5ell s Their nternal +rder6icrot*#*les 6aintain Cell Shape and 6ediate ntracell*lar Transport

6icrofila!ents enerate Contracting %orces and Sta#ili&e the

Cell S*rface

nter!ediate %ila!ents 7ary !ong Different Cell Types

2.0 The Cell Cycle and ts Control

Chro!oso!es re D*plicated D*ring S Phase

Chro!oso!e D*plicates re Split #et$een Da*ghter Cells D*ring 6itosis

The Cell Cytoplas! s Diided D*ring Cyto3inesis

The Cyclic ctiity of a Protein Co!ple4 Controls the Cell Cycle2. Cell 6e!#ranes

2.8 Cell*lar 6oe!ent

2.9 Cell :*nctions in Epithelia and 6esenchy!e2.; Cell*lar Signaling

ntercell*lar Signals 7ary $ith ,espect to Distance Speed of

ction and Co!ple4ity

6e!#rane ,eceptors nitiate Different Signaling Path$ays

denylate Cyclase enerates c6P as a Second 6essenger

Phospholipase C


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2 Kalthoff / Second Edition Contents

The ,TK


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$A%TE& / -A*ETO-E!E"I"

-.1 The Discoery of the 6a!!alian Egg

-.2 The er! "ine Concept and the D*al +rigin of onads

-.- 6eiosis'o!ologo*s Chro!oso!es re Separated d*ring 6eiosis

The Ti!ing of 6eiosis Differs #et$een 6ales and %e!ales6eiosis Pro!otes enetic 7ariation 'elps to Esta#lish

'o!o&ygo*s 6*tant lleles and Eli!inates Bad enes

-.0 Sper!atogenesis

6ale er! Cells Deelop in Se!inifero*s T*#*les

Sper!iogenesis

Sper!atogonia Behae as Ste! Cells

-. +ogenesis

+ocytes re S*pplied $ith "arge !o*nts of ,>

?ol3 Proteins for +ocytes re Synthesi&ed in the "ier or %at Body

The +arian *tono!y !poses Polarity on the +ocyte

6at*ration Processes Prepare the +ocyte for +*lation and

%ertili&ationEggs re Protected #y Different Types of Enelopes

@*ethod .' Autoradiora0hy

$A%TE&1 / FE&TILI2ATIO!

0.1 nteractions Before Sper!


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4.6 The Principle of Synergistic 6echanis!s

0.9 Parthenogenesis

3*ethod 1. I44unostainin

$A%TE&5 / LEA)A-E

.1 ?ol3 Distri#*tion and Cleaage Pattern

.2 Cleaage Patterns of ,epresentatie ni!als

Sea Archins 'ae solecithal Eggs and Andergo 'olo#lastic

Cleaage

!phi#ians 'ae 6esolecithal Eggs #*t Still Cleae

'olo#lastically

Snails 'ae solecithal Eggs and %ollo$ a Spiral Cleaage

Pattern

The scidian Cleaage Pattern s Bilaterally Sy!!etrical

6a!!alian Eggs Sho$ ,otational Cleaage

Eggs 5ith 7aria#le Cleaage Sho$ ,eg*latie Deelop!ent

Birds ,eptiles and 6any %ishes 'ae Telolecithal Eggs andAndergo Discoidal Cleaage

nsects 'ae Centrolecithal Eggs and Andergo S*perficialCleaage

.- Spatial Control of Cleaage Positioning and +rientation of 6itotic Spindles

ctin and 6yosin %or! the Contractile ,ing in Cyto3inesis

The 6itotic Spindle 4is Deter!ines the +rientation of theCleaage Plane

6echanical Constraints 6ay +rient 6itotic Spindles

Centroso!es +rgani&e 6itotic Spindles in ,eg*lar 5ays

D*ring Cleaage

Specific Sites in the Egg Corte4 ttract and nchor

Centroso!es6aternal ene Prod*cts 6ay +rient 6itotic Spindles

.0 The Ti!ing of Cleaage Diisions

The Cell Cycle Slo$s Do$n d*ring the 6id#last*la Transition

(6BT)

Stage< and ,egion< Specific ene ctiities 6od*late the

Basic Cell Cycle fter 6BT

The >*cleocytoplas!ic ,atio 6ay Trigger 6BT ccording to

a Titration 6odel

$A%TE&6 / ELLFATE, %OTE!(, A!DDETE&*I!ATIO!

8.1 %ate 6apping

8.2 Conal nalysis

8.- Potency of E!#ryonic Cells

8.0 Deter!ination of E!#ryonic Cells

Cell Deter!ination s Discoered Thro*gh +perational Criteria

DrosophilaBlastoder! Cells re Deter!ined to %or!

Str*ct*res $ithin a Single Seg!ent

Drosophila Cells re Born $ith a Bias that is 'oned #y


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S*#se*ent nteractions

Prospectie >e*ral Plate Cells of !phi#ians re Deter!ined

D*ring astr*lation

6o*se E!#ryonic Cells re >ot Deter!ined Antil the

Blastocyst Stage8. Properties of the Deter!ined State

Deter!ination s a Step$ise Process of nstr*ction andCo!!it!ent

Cell Deter!ination +cc*rs as Part of E!#ryonic Pattern

%or!ation

The Deter!ined State s (l!ost) Sta#ly Passed +n d*ring

6itosis

8.8 ,eg*lation in Deelop!ent

3*ethod 6.' La7elin ells 7y "o4atic rossover

$A%TE&8 / -E!O*IE9UI)ALE!EA!DT$E(%TO%LA"*IE!)I&O!*E!T

9.1 Theories of Cell Differentiation

9.2 +#seration on Cells

Cells 6ay Carry +*t Different %*nctions at Different Ti!es

Cells Change Their Differentiated State D*ring ,egeneration9.- +#serations on Chro!oso!es

Different Cells fro! the Sa!e ndiid*al Sho$ the Sa!e Set of

Chro!oso!es

Polytene Chro!oso!es Sho$ the Sa!e Banding Pattern inDifferent Tiss*es

Chro!oso!e Eli!ination s ssociated $ith the Dichoto!y

#et$een er! "ine and So!atic Cells

9.0 6olec*lar Data on eno!ic E*ialence9. Totipotency of Differentiated Plant Cells

9.8 Totipotency of >*clei fro! E!#ryonic ni!al Cells

>e$t Blasto!eres Deelop >or!ally after Delayed >*cleation

>*clei fro! E!#ryonic Cells re Still Totipotent

9.9 Pl*ripotency of >*clei fro! Differentiated ni!al Cells

>*clei fro! +lder Donor Cells Sho$ a Decreasing #ility to

Pro!ote the Deelop!ent of a >e$ +rganis!

>*clei fro! 6at*re Cells re Anprepared for the %ast 6itotic

Cycles of Early %rog E!#ryos

So!e Differentiated Cells Contain 'ighly Pl*ripotent >*clei6a!!als Can Be Cloned #y %*sing %etal or d*lts Cells $ith

En*cleated Eggs>*clear Transfer E4peri!ents $ith 6a!!als 'ae !portant

pplications in Science gric*lt*re and 6edicine5ill '*!ans #e Cloned in the %*t*reF

9.; Control of >*clear ctiities #y the Cytoplas!ic Eniron!ent

ene E4pression Changes *pon Transplantation of >*clei to

>e$ Cytoplas!ic Eniron!ents

Cell %*sion E4poses >*clei to >e$ Cytoplas!ic Signals

9.= Concl*sion and +*tloo3


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$A%TE&: / LOALI2ED(TO%LA"*IDETE&*I!A!T"

8.1 The Principle of Cytoplas!ic "ocali&ation

;.2 Polar "o#e %or!ation as a 6eans of Cytoplas!ic "ocali&ation

;.- er! Cell Deter!inants in nsects Eggs,esc*e E4peri!ents Can ,estore Defectie E!#ryos to

>or!al Deelop!ent

'eterotopic Transplantation Tests Cytoplas!ic Deter!inants

for ctiity in #nor!al "ocations

"ocali&ation of Polar ran*les ,e*ires ,> Transport long

6icrot*#*les

;.0 Bicoid !,> in Drosophila Eggs

;. 6yoplas! in scidian Eggs

Cytoplas!ic Co!ponents of scidian Eggs re Segregated

Apon %ertili&ation

6yoplas! is >ecessary and S*fficient for Tail 6*scle

%or!ation6yoplas! Segregation noles a Plas!a 6e!#rane "a!ina

6yoplas! is ssociated $ith "ocali&ed 6aternal ,>s;.8 Cytoplas!ic "ocali&ation at danced E!#ryonic Stages

;.9 Bioassays for "ocali&ed Cytoplas!ic Deter!inants

8.8 The Principle of Defa*lt Progra!s

;.= Properties of "ocali&ed Cytoplas!ic Deter!inants

Cytoplas!ic Deter!inants Control Certain Cell "ineages or

Entire Body ,egions

"ocali&ed Cytoplas!ic Deter!inants 6ay Be ctiating or

nhi#itory

Cytoplas!ic "ocali&ation 6ay or 6ay >ot Be ssociated $ith

7isi#le 6ar3ers6ost "ocali&ed Cytoplas!ic Deter!inants re 6aternal

!,>s

Cytoplas!ic +rgani&ation +cc*rs at Different Stages of

Deelop!ent Asing 7ario*s Cell*lar 6echanis!s

$A%TE&; / A


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11.1 Cell ggregation St*dies in Vitro

Cells fro! Different Tiss*es dhere to +ne nother Selectiely

Tiss*es %or! 'ierarchies of dhesieness

11.2 Cell dhesion 6olec*les!!*noglo#*linli3e C6s 6ay llo$ or 'inder Cell

dhesionCadherins 6ediate C2 H Dependent Cell dhesion

"ectins Bind 'eterot*pically to S*gar ,esid*es

11.- EC6 6olec*les and Their Cell*lar ,eceptors

lycosa!inopglycans and Proteoglycans %or! and

!orpho*s 'ydrophilic ro*nd S*#stance

%i#ro*s lycoproteins 6a3e Ap the Dyna!ic 6esh$or3 of

the EC6

ntegrins 6ediate Cell dhesion to EC6 6olec*les

11.0 The ,ole of Cell and S*#strate dhesion 6olec*les in 6orphogenesis

C6 E4pression is Correlated $ith Cell %ates

Cell dhesieness Changes d*ring Sea Archin astr*lation

C6s %acilitate the %or!ation of Cell :*nctions%i#ro*s EC6 Co!ponents Proide Contact *idance to Cells

!phi#ian astr*lation ,e*ires %i#ronectin on the nner

S*rface of the Blastocoel ,oof

11. 6orphoreg*latory ,oles of Cell and S*#strate dhesion 6olec*lesC6 and S6 enes re Controlled #y Selector enes

Cell


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12.0 6echanis!s of >e*ral nd*ction

There re T$o Signaling Path$aysJPlanar and 7erticalJfor

>e*ral nd*ction

Planar nd*ction Plays a 6aIor ,ole in Xenopus E!#ryos

>e*ral nd*ction is a 6*ltistep Process12. 4is nd*ction #y Disinhi#ition

Dorsal Deelop!ent +cc*rs as a Defa*lt Progra! in XenopusSpe!anns +rgani&er nactiates a 7entrali&ing Signal

Basic *estions are Still Anresoled

$A%TE&' / ETODE&*ALO&-A!"

1-.1 >e*ral T*#e

>ero*s Tiss*e Consists of >e*rons and lial Cells

The Spinal Cord s Patterned #y Signal fro! dIacent Tiss*es

The Basic +rgani&ation of the Spinal Cord s 6odified in the

Brain

The Peripheral >ero*s Syste! is of Dierse +rigin1-.2 >e*ral Crest

>C Cells rise at the Bo*ndary Bet$een >e*ral Plate and

Epider!is

>C Cells 'ae Different 6igration ,o*tes and a 5ide ,angeof %ates

The Strategy of Clonal nalysis Sho$s that >C Cells re a

'eterogeneo*s Pop*lation of Pl*ripotent Cells

The Strategies of 'eterotopic and 'eterochronic

Transplantation ,eeal Spatial and Te!poral ,estrictions to >C Cell

6igration

E4tracell*lar 6atri4 ffects the Deter!ination of >C Cells

,egionC CellDeter!ination

1-.- Ectoder!al Placodes

The +tic Placode %or!s the nner Ear

The "ens Placode Deelops Together $ith the ,etina

>asal Placodes %or! +lfactory Sensory Epithelia

1-.0 Epider!is

$A%TE&'1 / E!DODE&*ALA!D*E"ODE&*ALO&-A!"

10.1 Endoder!al Deriaties

The E!#ryonic Pharyn4 Contains a Series of rchesE!#ryos Pass thro*gh a Phylotypic Stage after +rganogenesis

The Endoder! "ines the nside of the ntestine and ts

ppendages

10.2 4ial and Para4ial 6esoder!

4ial 6esoder! %or!s the Prechordal Plate and the

>otochord

Para4ial 6esoder! %or!s the Preso!itic Plates and So!ites


10/20


So!ites re Patterned #y Signal fro! S*rro*nding +rgan

,*di!ents

10.- Connectie Tiss*e and S3eletal 6*scle

Connectie Tiss*e Contains "arge !o*nts of E4tracell*lar

6atri4S3eletal 6*scle %i#ers rise Thro*gh Cell %*sion

10.0 nter!ediate 6esoder!14.5 The Principle of ,eciprocal nteraction

10.8 "ateral Plates

The "ateral Plates S*rro*nd the Coelo!ic Caities

The Cardioasc*lar Syste! Deelops fro! 7ario*s

6esoder!al Prec*rsors

Cardioasc*lar Syste! Deelop!ent ,ecapit*lates the

Phylotopic Stage

S!ooth 6*scle Consists of Single Cells

10.9 E4trae!#ryonic 6e!#ranes

The !nion and Chorion re %or!ed #y "ayers of Ectoder!

and 6esoder!

The ?ol3 Sac and llantois re %or!ed #y "ayers ofEndoder! and 6esoder!

The 6a!!alian Placenta s %or!ed fro! the E!#ryonicTropho#last and the Aterine Endo!etri*!

$A%TE&'5 / T$EU"EOF*UTA!T"A!DT&A!"-E!IO&-A!I"*"I!T$EA!AL("I"OF

DE)ELO%*E!T

1.1 The 'istorical Separation of enetics fro! Deelop!ental Biology

1.2 6odern enetic nalysis of Deelop!ent

6*tants ,eeal the 'idden L"ogicM of E!#ryonic Deelop!ent

Drosophila6*tants llo$ the nalysis of a Co!ple4 BodyPattern

Caenorhabditis elegans6*tants Ancoer ene ctiities

Controlling the Cell "ineages

enetic nalysis of the 6o*seMus musculusAses E!#ryonic

Ste! Cells

The e#ra %ishDanio reriois enetically Tracta#le and

S*ita#le for Cell Transplantation

enetic nalysis ,eeals Si!ilarity of Plant and ni!alDeelop!ent

1.- D> Cloning and Se*encing

D> Can Be ,eplicated Transcri#ed and ,eerse*cleic cid 'y#ridi&ation llo$s the Detection of Specific

>*cleotide Se*ences

D> Can Be C*t and Spliced En&y!atically

D> Se*encing 6ay ,eeal the Bioche!ical ctiity of a

ene Prod*ct

1.0 Transfection and enetic Transfor!ation of Cells

15.5 The Strategies of ene +ere4pression Do!inant nterference and ene Knoc3o*t.


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1.8 er! "ine Transfor!ation

The enetic Transfor!ation ofDrosophilaAtili&es

Transposa#le D> Ele!ents

6a!!als Can Be Transfor!ed #y nIecting Transgenes

Directly into an Egg Pron*cle*sTransgenic 6ice Can Be ,aised fro! Transfor!ed E!#ryonic

Ste! CellsD> nsertion Can Be Ased for 6*tagenesis and Pro!oter

Trapping

Transgenic +rganis!s 'ae 6any Ases in Basic and pplied

Science

3 *ethods '5.' The -eneration and *aintenance of *utants

3 *ethods '5.+ "aturation *utaenesis "creens

3 *ethods '5. !orthern Blottin and "outhern Blottin

3 *ethods '5.1 &eco47inant D!A Techni?ues

$A%TE&'6 / T&A!"&I%TIO!ALO!T&OL

16.1 The Principle of Differential ene E4pression

18.2 Eidence for Transcriptional Control18.- D> Se*ences Controlling Transcription

,eg*latory D> Se*ences re St*died $ith %*sion enes

Pro!oters and Enhancers re ,eg*latory D> ,egions $ith

Different Properties18.0 Transcription %actors and Their ,ole in Deelop!ent

eneral Transcription %actors Bind to ll Pro!oters

Transcriptional ctiators and ,epressors ssociate $ith

,estricted Sets of enes and +cc*r +nly in Certain Cells

Transcriptional ctiators 'ae 'ighly Consered D>

Binding Do!ainsThe #icoid Protein cts as a Transcriptional ctiator on the

hunchback+ene in theDrosophilaE!#ryo

The ctiity of Transcription %actors The!seles 6ay Be

,eg*lated

18. Chro!atin Str*ct*re and Transcription

'eterchro!atic Chro!oso!e ,egions re >ot Transcri#ed

P*ffs in Polytene Chro!oso!e re ctiely Transcri#ed

D> in Transcri#ed Chro!atin s Sensitie to D>ase Digestion

Transcriptional Control Depends on 'istone cetylation and

Deacetylation

18.8 Transcriptional Control and Cell Deter!inationCo!#inatorial ction of Transcription %actors E4plains the

Step$ise Process of Cell Deter!ination

Cell Deter!ination 6ay Be Based on Bista#le Control Circ*it

of S$itch enes

Drosophila'o!eotic enes Sho$ S$itch ene

Characteristics

D> 6ethylation 6aintains Patterns of ene E4pression

3 *ethod '6.'In Situ$y7ridi@ation


12/20


$A%TE&'8 / &!A %&OE""I!-

19.1 Posttranscriptional 6odifications of Pre19.2 Control of Deelop!ent #y lternatie Splicing

Cascade of lternatie Splicing Steps Controls Se4Deelop!ent inDrosophila

lternatie Splicing of Calcitonin and >e*ropeptide !,> s

,eg*lated #y Bloc3age of the CalcitoninP Transport fro! >*cle*s to Cytoplas!

>*clear Pore Co!ple4es re Controlled ates for the

Transport of ,>Ps to the Cytoplas!

E4peri!ents $ith Cloned cD>s ndicate Differential !,>P

,etention

19.0 6essenger ,> Degradation

The 'alf s Stored in S*#ri#oso!al ,>P Particles

#efore Translation

6essenger ,> 6ay Be Ear!ar3ed for Storage #y ts

AT, or - A,T Se*ence

1;. Posttranslational Polypeptide 6odifications


13/20


Polypeptides re Directed to Different Cell*lar Destinations

Polypeptides 6ay Andergo Seeral Posttranslational

6odifications

Protein Degradation s Differentially Controlled

$A%TE&'; / -E!ETIA!D%A&A-E!ETII!FO&*ATIO!

19.1 The Principle of enetic and Paragenetic nfor!ation

1=.2 Self


14/20


The Sa!e Aniersal Ste! Cell %or!s ll Types of Blood Cells

as $ell as Endothelial Cells

Progenitor Cell Co!!it!ent 6ay Depend on Sta#le Control

Circ*its of enes and Transcriptional %actors

Blood Cell Deelop!ent Depends on Colony


15/20


and #y "ocal nteractions

$A%TE&++ / -E!ETIA!D*OLEULA&A!AL("I"OF%ATTE&!FO&*ATIO!I!T$EDROSOPHILA

E*B&(O

22.1 ,eie$ ofDrosophila+ogenesis and E!#ryogenesis22.2 Cascade of Deelop!ental ene ,eg*lation

22.- 6aternal enes ffecting the nteroposterior Body Pattern

The nteroposterior and Dorsoentral 4es +riginate %ro!

the Sa!e Signal

The nterior ro*p enerates and *tono!o*s Signal

The Posterior ro*p cts #y Depression

6any Posterioret$or3

22.8 The Dorsoentral Body Pattern

22.9 The Co!part!ent 'ypothesis

Co!part!ents re Controlled #y the ctiities of Specific

Selector ene ctiities

The Ultrabithorax+E4pression Do!ain Coincides $ith aCo!part!ent

Engrailed+Controls the nteroposterior Co!part!ent

Bo*ndary

Apterous+Controls the Dorsoentral Co!part!ent Bo*ndary22.; ppendage %or!ation and Patterning

Co!part!ent nteractions across Bo*ndaries +rgani&e Pattern%or!ation $ithin Co!part!ents

Decapentaplegic Protein %or!s a 6orphogen radient in the

5ing !aginal Disc

22.= Patterning enes in +ther nsects

$A%TE&+ / T$E&OLEOFHOX-E!E"I!)E&TEB&ATEDE)ELO%*E!T


16/20


23.1 Strategies for dentifying Patterning enes in 7erte#rates

enes re Cloned on the Basis of Their Chro!oso!al 6ap

PositionPro!oter Trapping dentifies Patterning enes

7erte#rate enes Can Be solated $ith 6olec*lar Pro#es fro!

Drosophilaenes

2-.2 'o!eotic enes andox enesThe 'o!eodo!ain is a Cooperatie D>


17/20


Pattern %or!ation in Plant E!#ryos noles 2 to G Specific

ene %*nctions

ro*ps of enes Control Distinct Patterning Eents

20.- enetic Control of %lo$er Deelop!ent

%loral nd*ction enes Control the %or!ation of annflorescence

6eriste! dentity enes Pro!ote the Transition fro! %loral6eriste!s to nflorescence 6eriste!s

20.0 The ,ole of 'o!eotic enes in %lo$er Patterning

Arabidopsis'as 'o!eotic enes

Three Classes of 'o!eotic enes Deter!ine the

6orphological Characters of %o*r %lo$er 5horls

Do*#le and Triple 'o!eotic Phenotypes Confir! the enetic

BC 6odel

'o!eotic enes are Controlled #y ,eg*lator enes 6*t*al

nteractions and %eed#ac3 "oops

+ther Plants 'ae Patterning enes Si!ilar to Those of

Arabidopsis

Antirrhinum'as enes Controlling +rgan 7ariations 5ithinthe Sa!e 5horl

20. 6olec*lar nalysis of 'o!eotic Plant enes

The agamous+ ene Encodes a Transcription %actor

enetic and 6olec*lar St*dies ,eeal +rthologo*s enes#et$eenArabidopsis andAntirrhinum

The Kno$n Plant 'o!eotic enes 'ae a 6DS Bo4

nstead of a 'o!eo#o4

$A%TE&+5 / Eor!al Deelop!ent

'er!aphrodites and 6ales

%ertili&ation Cleaage and 4is %or!ation

%o*nder Cells

astr*lation +rganogenesis and 'istogenesis

"aral Deelop!ent2.2 "ocali&ation and nd*ction D*ring Early Cleaage

The %irst Cleaage enerates Blasto!eres $ith Different

Potentials

P ran*les re Segregated into er! "ine CellsPar enes ffect Cytoplas!ic "ocali&ation

Three 6aternal Effect enes enerate a "ocali&ed ctiity thatPro!otes E6S dentity

Deter!inant ctiity for the E6S Blasto!ere

Deter!ination of nterior Pharyngeal 6*scle Cells ,e*ires

6*ltiple nd*ctie nteractions

P2 Polari&es E6S to %or! Ane*al Da*ghter Cells

2.- 'eterochronic enes

6*tations in the lin!"#+ene are 'eterochronic

The lin!"#+ene Encodes a >*clear Protein That %or!s a


18/20


Te!poral Concentration radient

The lin!#+ene Encodes S!all ,eg*latory ,>s $ith

ntisense Co!ple!entarity to lin

Do$n


19/20


So!atic onadal Cells

$A%TE&+8 / $O&*O!ALO!T&OLOFDE)ELO%*E!T

29.1 eneral spects of 'or!one ction

29.2 'or!onal Control of Se4 Differentiation in 6a!!alsThe Synthetic Path$ay for 6ale and %e!ale Se4 'or!ones

re nterconnected

The enital D*cts Deelop fro! Parallel Prec*rsors the

5olffian and 6llerian D*cts

6ale and %e!ale E4ternal enitalia Deelop fro! the Sa!e

E!#ryonic Pri!ordia

29.- 'or!onal Control of Brain Deelop!ent and Behaior in 7erte#rates

ndrogen and Estrogen ,eceptors re Distri#*ted Differently

in the Brain

ndrogens Ca*se Seasonal and Se4*ally Di!orphic Changes in

the Brains of Birds

Prenatal E4pos*re to Se4 'or!ones ffects d*lt,eprod*ctie Behaior and Brain nato!y in 6a!!als

29.0 'or!onal Control of nsect 6eta!orphosis

nsect 6olting P*pation and 6eta!orphosis re Controlled

#y 'or!onesP*ffs in Polytene Chro!oso!es ,eeal enes ,esponsie to

Ecdysone

6*ltiple Ecdysone ,eceptors 'ae Tiss*e


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2;.0 ro$th %actors

>ere ro$th %actor Pro!otes the ro$th and Differentiation

of Certain >e*rons

+ther ro$th %actors ffect 6*ltiple Target Cells in a 7ariety

of 5aysThe Effects of a ro$th %actor 6ay Depend on the Presence

of +ther ro$th %actors2;. 6echanical Control of Cell S*rial and Diision

2;.8 Cell Cycle Control

2;.9 T*!or*!#er

$A%TE&+; / "E!E"E!E

2=.1 Statistical Definition of Senescence

2=.2 Eol*tionary Theory of Senescence

The 6*tation cc*!*lation 'ypothesis %oc*ses on the ge

Distri#*tion in >at*ral Pop*lations

Senescence Can Be scri#ed to ntagonistic Pleiotropy

ccording to the Disposa#le So!a 'ypothesis Eeryndiid*al is the Te!porary Carrier of 'is/'er er! "ine

2=.- 7ario*s Physiological Ca*ses of Senescence

2=.0 Cell*lar Senescence and Telo!erase

So!atic 6a!!alian Cells Sho$ a "i!ited Capacity for

Proliferation

Chro!oso!al Ends re Protected #y Telo!eresTelo!erase ctiity Co*nteracts Cell*lar Senescence

"oss of Telo!erase ctiity is a Safeg*ard gainst Cancer

2=. Da!age fro! +4idants and +rganis!ic Senescence

+4idatie Phosphorylation enerates ggressie +4idants asnter!ediates

+4idants Da!age Cell*lar D> "ipids and ProteinsS*pero4ide ,adical Sti!*lates Cell Diision and ro$th

ntio4idatie gents Delay Senescence

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