Lewin Benjamin (ed.) Genes IX

daughter

chromosomes

to separate;

each of

them has

already

been

partially

replicated

by

the new replication

forks

(which

now

are the

only replication

forks).

These

forks

continue

to

advance.

At the

point

of

division,

the two

partially

replicated

chromosomes

segregate.

This recre-

ates the

point

at which

we

started. The

single

replication

fork

becomes

"old,"

it terminates

at

l5

minutes, and 20

minutes

later

there is a

divi-

sion. We

see that the initiation

event

occurs l,Zs

cell

cycles before the

division

event

with which

it is

associated.

The

general

principle

of

the link

between

initiation

and the

cell cycle

is that,

as cells

grow

more rapidly

(the

cycle is shorter),

the

initiarion

event occurs

an increasing

number

of cycles

before the related

division.

There

are

corre-

spondingly

more

chromosomes

in the individ-

ual bacterium. This

relationship

can be viewed

as the

cell's response

to its inability

to reduce

the

periods

of C and D

to keep

pace

with the

shorter

cycle.

@

The

Septum

Divides

a

Bacterium

into

Progeny

That

Each

Contain

a Chromosome

Septum

formation

is initiated

at the annu[us,

which is

a ring around

the cetl where

the

structure

of the envelope is

a[tered.

New

annuti are initiated

at 50%

of the distance

from the

septum to each

end ofthe

bacterium.

When the bacterium

divjdes,

each daughter has

an

annulus at the mid-center

position.

Septation

starts when the

ce[[ reaches

a

fixed

te n

gth.

The

septum consjsts

of the same

peptidogLycans

that comprise the

bacterjaI envelope.

Chromosome

segregation

in

bacteria is espe-

cially interesting

because

the DNA itself

is

involved

in the mechanism

for

partition. (This

contrasts with eukaryotic

cells,

in which

segre-

gation

is

achieved by the

complex apparatus

of

mitosis.)

The bacterial

apparatus is

quite

accu-

rate; however,

anucleate cells form <0.03%

of

a bacterial

population.

The division

of a bacterium into

two daugh-

ter cells is accomplished

by

the formation

of a

septum, a

structure that forms

in the center

of

the

cell as an

invagination

from the

surround-

ing envelope. The

septum forms

an impenetra-

ble barrier

between the two

parts

of

the cell and

provides

the site at which the two daughter cells

eventually

separate entirely.

TWo related

ques-

tions address

the

role

of

the septum in division:

What

determines the

location at

which

it forms,

and

what ensures that the daughter

chromo-

somes lie on opposite sides of

it?

The formation

of the septum

is

preceded

by the organization of the

periseptal

annu-

lus.

This is observed as a

zone in E. coli or Sal-

monella typhimurium, for which the structure of

the envelope is

altered so

that the inner mem-

brane is connected more closely to

the

cell

wall

and outer membrane layer. As

its name

sug-

gests,

the annulus extends

around the cell.

i'ir.i-

lirii

r

i'

summarizes

its development.

The

annulus starts at a

central

position

in

a

new

cell. As the cell

grows,

two events occur:

A septum forms at the mid-cell

position

defined

by the annulus,

and

new annuli form on either

side of the initial annulus.

These new annuli

are

displaced

from

the

center and move along

the cell to

positions

at one

quarter

and three

quarters

of the cell

length.

These

will become

the mid-cell

positions

after

the next division.

The displacement of the

periseptal

annulus to

i::i::rtl

l t

i

r

Duplicationand

dispLacementofthe

perisep-

tal annutus

give

rise

to

the formation of

a septum that

divides the cetl..

I

t

I

Cell starts with

annulus

at midcenter

New annuli are

generated

New annuli

move

in

polar

direction

New annuli stop

movemenl

Central annulus

develops

into seplum

Side

view

shows that

annulus extends around

circumlerence of cell

Cross-section shows

that

annulus

connects

membranes

17.3 The

Sentum

Divides

a

Bacterium

into Proqenv

That Each Contain a Chromosome

4tt

by

the simple extension

of a

polymeric

struc-

ture. Another enzyme

is responsible for

gener-

ating

the

peptidoglycan in the

septum

(see

Section

17.5,FtsZIs Necessary

for

Septum

For-

mation). The septum

initially forms as a double

layer of

peptidoglycan,

and the

protein

EnvA

is

required to split the covalent

links between the

layers

so that

the daughter cells may separate.

The behavior of

the

periseptal

annulus sug-

gests

that

the mechanism

for

measuring

posi-

tion is associated with

the cell envelope. It is

plausible

to suppose that the envelope could

also be used

to ensure segregation of the chro-

mosomes.

A

direct

link between DNA and the

membrane could account

for

segregation.

If

daughter chromosomes

are attached to the

membrane, they could be

physically

separated

when the septum

forms.

Fl+#F{

1}"s+ shows that

the formation of a septum could segregate the

chromosomes

into the different daughter cells

if the origins are connected

to

sites

that lie

on

either

side of the

periseptal

annulus.

i:i.:i:i:::l

':

..-

Attachment of bacteriaI DNA to the mem-

brane could

provide

a mechanism for segregation.

the correct

position

may be the crucial event

that ensures the division of the cell into daugh-

ters

of equal size.

(The

mechanism

of

move-

ment is

unknown.) Septation begins when the

cell reaches a fixed length

(2L),

and the distance

between the new annuli

is

always L. We do

not

know how the cell measures length, but the

relevant

parameter

appears to

be

linear distance

as such

(not

area

or

volume).

The

septum consists of the same compo-

nents

as the cell envelope: There is a rigid layer

of

peptidoglycan

in

the

periplasm,

between

the inner

and outer membranes. The

peptido-

glycan

is made by

polymerization

of tri- or

pentapeptide-disaccharide

units in a reaction

involving connections

between both types of

subunit

(transpeptidation

and transglycosyla-

tion). The rodlike

shape of the bacterium

is

maintained

by a

pair

of activities, PBP2 and

RodA. They

are interacting

proteins

and are

coded

by the same operon. RodA is a member

of the

SEDS

family

(SEDS

stands for

shape,

elon-

gation,

division, and

sporulation) that is

pres-

ent in

all bacteria that have a

peptidoglycan

cell

wall. Each SEDS

protein

functions

together with

a

specific transpeptidase,

which catalyzes the

formation

of the crosslinks in

the

peptidoglycan.

PBP2

(penicillin-binding

protein

2) is the

transpeptidase

that interacts with RodA. Muta-

tions in the

gene

for

either

protein

cause the

bacterium

to Iose its extended

shape and become

round. This

demonstrates the important

prin-

ciple that shape and rigidity

can be determined

CHAPTER 17 Bacterial

Reptication

Is Connected to the

Ce[[ Cycte

Origins of

replicating

chromosomes

attached

to membrane

Daughter chromosomes

attached to envelope

Septum

grows

between chromosomes

Septum

divides cell

Chromosomes

distributed to

daughter cells

Mutations in Division

or Segregation

AfFect

Ce[[ Shape

o

jfs

mutants form long fitaments because the

septum

fails to form to divide the daughter

bacteria.

o

Minicelts form in mutants that

produce

too many

septa;

they are sma[[ and Lack DNA.

o

Anucteate cetts of

normal

size are

generated

by

partition

mutants

in

which the dupticate

chromosomes

faiI

to seoarate.

A difficulty in isolating mutants that affect

cell

division is that mutations

in

the critical functions

may

be

lethal

and/or

pleiotropic.

For example,

if formation of the annulus occurs at a site that

is essential for overall

growth

of the envelope,

it would be difficult to distinguish mutations

that specifically interfere

with

annulus

forma-

tion from those that

inhibit

envelope

growth

generally.

Most mutations in

the division appa-

ratus have been identified as

conditional

mutants

(whose

division is affected under non-

permissive

conditions; typically they

are tem-

perature

sensitive). Mutations

that affect cell

division or chromosome segregation

cause

striking

phenotypic

changes.

Ftr{itlfito"

l;.*

and

It"$ul{F.

i,r"ii

illustrate the opposite consequences

of

failure

in the division

Drocess

and failure in

segregation:

472

EI'

uoqeurol

urntdes

toJ &pssalaN

sI

zsll

g./I

.O1\^I

OIUI

IIJJ

JqI

q]UId

ol

slJrJlsuoJ dprrnb

uaql

pup

'salnuru

s I

JoJ

surPruar

'uorsr^rp

JelJe selnurru

E

01

I

IIal

aqt

Jo

JelueJ

aql ur surroJ

ll

'apdr

uorsr,rrp

lerrd;(t

e uI

'uorleuro;

runldas

ur dals 3ur1nu11-ater

aql sr Eurr-7

aql

Jo

uortpuroJ

aqJ

'g'11

arn8r4

Jo

snlnuue raluaJ-pru

eql

Jo

uorlrsod Jql ul

sa1

ll reqt

smoqs

I'lt

lunlH

'tu;t-7

aql

prlleJ

saurrlaruos

sr aJnlJnJls

aqJ

'atuareJrunJrrf,

aql

punoJp

Surr

e ur

pazr1pJol

saruof,aq

Zsld

'alpp[u

eql uI

t)lJtsuoJ

o1

sur8aq

pue

saleEuole

IIaJ

eql

sV

'urse1do1[r

aqt

lnoq8noJql

pezrlpJol

sI

ZSld

'a1rz(r

uorsrn1p

aqt ur ,{.1teg

'uorteruro;

runldas;o

a8els ,{.pea

ue

le

suorpunJ

Zsld

'UOI]EZIIEJOI

IIAqI JO

qnuue

pldasFad

aqr

yo

uorlerruoJ aql

DeJIp

Jleslr

lou

saop

lnq

'uorlpuroJ

urnldas

ro; salrs

3ur

-tslxaard;o

a8esn ro;

parrnbar

eJoJaJJql sl

ZStJ

'srsauaSoqdrou

pldas

01 rlnuup

ptdasrrad

aql

Jo

tualxeJpldsp

aqt ruor;

Suuhen sa8els

te tre

sluetnurZs{'sselu

IIaJ

l1un

rad

slue^a uopetdas

Jo

rJqrunu

paspaJJur

uB Sursner z(q sllatrurur

saJnpur

uorssardxaraAO'sluJruell;

alerauaB

puP

uorleurro;

runldas

polq

7sl

ur suortptnl\l

'uorsrlrp

ur

eloJ

lprluJ)

e sz(e1d

ys{

aua8 aq7

'sluauodnroc

1e1a1aqso$r

raqlo

ol

palleuuol

sr

11

'edolanua

lpueppq

aql

Jo

aprsul aq1 uo 6uu

e suloJ

leql

aspdlg

p

st

ZslJ

'sa1rs

6uqsxaard

1e

uorleuro;

unldas

rol

parrnbar

t

7s{

1o

pnpord

aq1

uoqeurol

unldas loJ

firessaraN

sI

Zsll

'(uorleBa.rBes

auosoruoJqJ

ur

enrlJeyap are l(aqt

JSneJaq

parueu)

sluptnru

(uoltltred)

wd trqpasneJ

sr

addlouaqd srql'para1

-lPun

sr JZrs Jlar{l

'plurou

sr uorlpruJoJ

urnldas

esneJaq

lnq

'auosourorq)

p

1re1

,{.aqr

'sllJ)rurur

a>lq

ltupJreqe

sr

uorl

-BBarEas

uJqM ruJoJ sllal

alBaIJnuV

'lpruJou,{.1er6o1oqdroru

sreadde asr,u

-rJqlo

lnq'vNC

s>l)PI

puP

ezrs

IIPUS

JJqIPJ e

sPq

llaJrultu

JqJ

'aurosour

-oJqJ

e s>lJpl sllal

ralq8nep ,lr.au aqt

Jo

auo

leql tlnseJ

aql

qtyu

'are1d

8uor.,lr

eql ur ro z(lluanbalJ

ool sJnJJo uorl

-pruJo;

urnldas

uaqm ruroJ

sllalfuf!\t

r

';1as1r

ssarord

uorsr^rp

Jql

q

all

tpql

spJJJp aldglmu ro

tJa;ap

p

,$puapr qJIqM'(uopeluaurellJ

elrlrsuas

-arnle.radrual

ro;

pauteu)

sluelnru s/u(q

paleldsrp

sr ad$ouaqd srqJ

'llar

aqt

Jo

et8ual

aqt Suop

patnqlrlslp

[gelnEar

(saurosoruorqr prrapeq)

sproalJnu eq1

I{tl1vr

'alnlJnJls

snolualueg; 3uo1

dra,r

p

Jo

slsrsuoJ

IIaJ

Jq1 snr{J

'rrrJoJ

lou

op

eldas

tnq-sauosoruoJq)

ralqEnep rraql

aleEarEas 01 anrrrfuo) uazla

pue-u.or8

o1

enurluoJ Puapeq ar{J

'papaJIeun

$ uoueJ

-r1da:

auosoruoJqJ

lnq'pJllqlqq

sr uoll

-euuo;

runldes uar{,lrrruoJ

guawal{8uo1

.

'Alrsranrug

olor{;

'eberrg

e1o5;o r{se1

-rnol

oloqd

'uaas

eq uel suolsr^lp

lPurouqP

puP

lPuilou

qloq

jluelnu

Olnw

eql

J0

qlal

sMoqs

plalJ

slql

'urpls

onlq ou o^eq sauosourorqr

6uqre1 slac ralqbnep 1an1q

u rpls soulosouro.l

ql qlu'A qlal'sl

lpJ

uorlebarEas eurosour

-orqr

uaqm sllal alpallnue

elereueb

4or

j g'LL

lgflgll

'r{lgsranruq

olor{y'e6errg

e1o5;o r\salnor soloqd

'slueurellJ

paleallnuqlnu

salera

-ua6

sernle.radual anrssu.uraduou

rapun uolsl

lp

llar Jo

aln

-ye1ryuod

wo17og'qlar ad&-pll4

:1auod do1

9'lI

lgngll

.UOIIJIJ]SUOJ

E UJOJ O1 JUPIqIUJIII

Jq} SJqSNd

ro sllnd rJqtrJ

pup

JIIeueSJo aqt

punoJe

SurJ

e sluJoJ

leql

ureloJd

lelela>lsol^J

p

Jo

JSn Jql

sr PrJpuoqJollru

pue

'slsPIdoJolqJ 'PrrJtJeq

Jo

uorsr^rp eql uUuJql

'JJnlpJI

uotutuo) aql

'uorl)rJls

-uoJ

e JleJauJB

ol JuprquJru Jql Surzaanbs

'a1aue3ro

Jql

Jo

eprstno

Jqt uorJ suorlJunJ srql

'ruseldolfu

rtlo.{re>lna

Jo

sJuprqruJur ruor}

sJIl

-rsel

Jlo

Surqrurd ur

pJAIoAur

sr

qlrqu

'utueu.{p

uratord Jqt

Jo

tueuel

e esn [aqt

'pealsul

'Zstd

alpq

lou

op.{1ensn

'erre1rpq

qtrm

ur8rro d-reuorl

-nlo^J

uP JrPqs oslP

qJrqM

'Plrpuorlfolll^l

'pJAJJSUoJ

uJJq a^eq ol L11e

-Jeue8

sureJs uorsrnrp roJ

snle-redde aqt

'stseld

-orolqJ

pue

prJelJeq

;o

sur8rro

,{reuortnloa,a

uoruruoJ Jrll

qlr.,r,r

lualsrsuoJ

'saua8

uorsrlrp

IeIJJtJeq

Jqt ol

pelelal

seua8 raqto

J^pq osle

s1se1doro1q3

'rseldorolqr

Jqt

Jo

lurodpnu

aql

le

Surr e 01 sSoloruoq

lueld

Jql

Jo

uorlezrlerol

Jql sMoqs

i:j'.:

-!

:iIii:tii:,

'slseldorolqr

ur

punoJ

sr

'AlLsranrul

a1e15 uebrqrr6l

'6uno&a1s6

aur

-laqle)

lo

fsalnor

so1oq4

'fipee1r

etou

lseldotolqr

aq1

Jo

ourlno eql sMoqs (1aued

ramol)

ebeurr

plag

tqbyq

aq1

'(1aued

do1)

lseldorolqr

aq11o

lurodprLu

aql

le

pezrtelol

ele

[aq1

1eq1

Moqs

Zzsll

pue

IZs]l

suLelord sLsdopqoty aq1

lsuLeEe

saLpoqque

qlm

eluelsalongounuul

lt

;.

i

:iHitl-l_l

a1rf1

11a1

aqt ot

pelrauuol

sI uoqelrtdau

lPuapPE

osle

pup

'erJelJpq

ur uoruuro) s1

t1

'uoneldas

;o

tueuoduoJ

letalalsotr{r

roleru eql sr

Zsl{

'JUPJqTUJUT

rJtno

Jqt 3ur11nd

pue

JuerquJur

Jauul eql

Surqsnd

snqt

'prpuur

,rtor8 ot

uer.r(l3oprl

-dad

aql sesneJ uJqt

^trlrDe asepqdadsuel aql

qJIqM

ur uorlpruJoJ unldas

roJ

IJporu

B slsa8

-3ns

srql'3ur,r

leldas

Jqt

olur

Isl{

Sutlerodrorut

ro; alqrsuodsJr sr

14sld

'ruseldt:ad

Jql

uI atIS

rr/.1e1er slr seq

leql

ulalord

punoq-JueJqruau

e

'(g

uralord Surpurq-urpluad

roJ

€dgd

pellpJ

osle)

aseprtdJdsuprt

e ro; sJpor

qJIqM

'Is{rlty*

uoJJdo ue

;o

lred

se

passardxa

sI

AS{',{1ue;

S11AS

eqt ot Sur8uolaq utatord e sI

q)Iq,lt.'MSld

sl

Surr

1e1das

eql otur

pJterodroJul

Jq 01 slueuod

-ruoJ

lspl

Jqt

Jo

Juo

'Juerqueur

aql

lJrJlsuor

or r(lrlrqe Jql J^pq o1

parunsard

sI

teql

xaldruor

uratordrtlnru e

Jo

stsrsuoJ

tI

'EuIr

pldas

aqt

pJIIeJ

sJrurlauros sr JJnlJnJls

Ieur}

JqJ

'sural

-ord

auerqruJursupJt

1e

are

Laql'pJteJodJorur

urrq

seq

vst{

JJue

rrpro

peurlep

e ur 8uu-7 aqt

uroI saua8 sy'raqto

IeJJAas

yo

snnpord aq1

'JuerquJru

Jq1

ot

tr

Suuluq ur sdeq;ad

pue

3ur:-7 Jqt SUIZITIq

-ets

ur seloJ Surddega,ro azr.eq

daqt

teqt

s1sJ33ns

srqJ

'luasqP

Jre

qloq

Jr

uJoJ

louue) lnq

'vslJ

ro

ydrT

reqlra

Jo

aJuasqe aqt

uI ruJo; uetBurt-7

eql'eueJquJur er{] r{]r^^

palPrJosse punoJ

ue{o

sr

tnq

'uralord

rrlosol.dr

p

sr

vslC

'euerqruJru

Jql

(]1

zslg

Sullull

ro} sueeur aqt saprrrord

11

'JuPrquJu

IerJJDeq

JJuur Jql

ur

pateJol

sr

lPql

uratord

JuprqurJru

pr8alur

ue sr

ydrT

'ZSl{

qlIM

.dltuapuadapur

pue,{prarrp

DeJJtul

'VSt{

pup

ydrT

'uorsr.Lrp

roJ

pepJeu

suralord raqto o,t41

'1ood

rtuseldoldr e

qllM

slrunqns;o a8ueqrxJ snonulluoJ

sI JJJqI

qJrqm

ur

'erntJnJls:nueuLp

e sr?urt-7 eqJ

'aJnl

-:nrls

3ur-r Jql olur sJJruouoru

ZSI{

}o

uollpzl

-raruo811o

aqr

uoddns

ol

pJSn

sr aSerrealr

419

pue

.{.1rnrlre JSedID spq

ZS1C

's11ar

ruo.{.re>lna

ur selnqnloJJ[u

Jo

uorleuJoJ aql alqueseJ

plnor

Surr

Jqt

Jo

,{.lqruasse

teqt

SurlsaSSns

'urlnqnl

salqussJJ

zsl{

Jo

eJnl)nJls eqJ

'uolsnoH

le

looqrs lPrr

-pa61

sexallo fi1rsre^run

'uqobrejl

Lueq1L1111o [se1no] oloqd

'llar-prur

aql

le

pazrlprol

sr

lr lpql

sMoqs

Zsll

lsuLe6e

r\poqLlue ue

qlrM

arualselonl1ounuul

l.a'!

$*f]=.a-lij

/r

ulldvHl ,r,

91?

uotleurquolau

lLloads-alrs

elrnbau

[e6

uorlebarba5

leuosoruo)tlJ

L' Ll

'rautp

P o1 auosorlolql

lPuolreq

aql

paila^u0r

seq

lua^a

uo4purqulotaj

pazrlereueb

p

Jr

sloruououl

alpallal oJ snurulal

ourosoruorqr

aql

leau aruenbes

labte1

e uo

slre rlalsfs

uorleurquotal

rgoeds-a1rs raf otlf

r

uor.lPurquolau

lqpads-alrs

alrnbag Aeyrl

u orJP6el

6a5

1

eur osoru

oJ

ql

'8urr

gurl,rg

aqt

Jo

uortpruroJ

roy

parrnbar

sr

qur141

Llsno

-trn3

'(ydt7

pup

ZSt{

sJpnlJur

qrlrlm)

3ulr

letdas

Jqt

Jo

uorteuro;

3ur.tro1e snqt []ruor^ Jql ur

Clulw

Jo

uoltf,e aqt sassarddns

uoltelnrrlnJle

sU

'uorlrsod

1e1das

aqt

le

Surr e sruJol

AurW

'uorlelualuelrJ

ur 3ur11nsar

'salrs

relod

sp

IIaM

se

IIJJ-prtu

sllqlqul

Eulw

Jo

rf,ursqe ro

o)urw

qrnu

oo1 lslleJrunu

Sunuro;'uorleldas

JleunurrJsrpur

sJsnel

'gulw

qJnu

ool ro

'c)urw

Jo

e)uJSqP

'lu!W

^q

salLs relod aql o]

pauL+uol

sr

uorJlp

osoqM loJrqrqur uorsr^rp

e sL

6/3u161

:palelar AIJSJaAUT JJP

guIW

puP

Q/)uIW

Jo

speJJJ aqJ

'uouetdes

IIaJ-prru

slturad

1nq

uoqel

-das

relod sluanard

1ana1

adr{t-p1,tr

JqJ

'furw

ol

q1:)ulw

Jo

orter rqt

'JroJJreql 'sI

atIS

(11ar-ptu)

radord aqt

le

uorteldas;o

tueunuJJtJp

JqJ

'suorteJol

rpoq

le

ruroJ ol

eldas Sutrltolle

'llaJ-pnu

te

se

IIaM

se salod Jql

te

uoltlqlqur

aql slJeJelunor

[urw

ssJJXJ;o eruasard aql

JSne)Jq

'sllJJruru

seJnpur

gulw

Jo

uolssJrdxe

-JJ^O

'uoulqrqur

uroJJ satrs

IIJJ-pnu

aql sDalord

AurW'qtmor8

prurou

Sur.rolsar os

'suot3ar

relod

rqt ot uoplqrqu

eqt saulJuoJ

Olulw

o1 alqe.red

-ruoJ

slJ^JI

tp

gurw

Jo

uotssardxg

'e1das

lnoqlptr

stueuelrJ 3uo1 se

rr,to.r8 s11ar 3ur11nsar eqJ

'uoIS

-l^lp

Jo

uorlrqrqur

pazlleraua8

p

sJsnpJ

'EulW

Jo

aruasard aql

ur UJAJ uorssa:dxaralo

Jo

'guIW

Jo

J)uJsqp JLI

I

ur

q)ulW

3o

uorssa.rdxl

'

(3urt-7

aql olur Surzr

-raru.dlod

urorJ

Zs1{

sluanard

qrtqM

')uIW

J}eA

-rtJe

ot

parrnbar

sl

11uIW

'Jolrqlqul

UoISIAIp e

r;lJo!eullupue)unalo

spnpord

aq1

','

-

:tt.

j

ur

pazrrPurruns

eI€

saloJ JIeqJ

'g-

pue

'Q-

'3utw'saua1

JJrql

Jo

slsrsuol

s\)olgunlt JrlI

'uotleldas

JlpelJnu

ueJ slupuurer

aseqt

pup panrJep

aram

Laqt

qrlqM

uorJ

llnuup

eql

Jo

slupuruJJ

urplal salod

eql sdeqra4

'1eql

JroJaq

uorsrlrp eqt

uorJ runldas aql stuasardar

alod raqlo Jr{l

luolsll1p snotnard

aql;o unldas

Jql

uorJ

peuroJ

se,r,r

alod aug

'sa1od

Jql

qllM

pue

JJlueJ-pru

1e

snlnuue

aqt

qll^^ qloq

pJlerJ

-ossp

selrs uorleldas

prlualod

seq

[a)

uJoqMau

p

teql

salldur

srqt

'€'L

I

arn8rg ur

palrldap

sluJle

aqt

Jo

sruJel u1

'sa1od

aql

le

uoltetdas ssarddns

ot sr

snrol

gurut

ad1't-plrm

aql

Jo

JIor aqt

rpql

pue

'sa1od

Jqt

te

ro

IIJJ-pItu

tp

Jaqlla uollpru

-roy

runldas etBrtrur

o1 ,{.1rgqe aql

sassassod

1ar

aqt

teqt

srsaSSns sIqJ

'llrl-plru

1e

(yo perlsul

ro) se

11aur,r

se salod eql

t€

rnlJo

o1 uorleldas 3ut

-Molle

Lq slarrunu sateraua8

guttu

lo

uortalJp

:gutw

snJol Jr{l

uI serl uollelnlu

IIJJIUIU

IeuI

-3rro

aql'sluptnru

IIJJIUITU,{q

papnord

sI runl

-das

aqt

Jo

uorlezllpf,ol

aqt

tnoqe

uolleurloJul

'sa1od

aq1

1e

6uttttto; tuot;

lu!y'l

Jo

sbuu

leuoqLppe

sluenerd

1nq

'6uu

taluar

-pru

e sMolle

6/1u114

'suotlerluetuol

leulou

lV

'DUl'.1

e uloJ ol

elqe st

lur6

oraqM

suuoJ unldas aq1

'0'lulw/lult/,1

Jo

otlel aql

Aq

peururalap

sr eldas

jo

uorlplol

puP

loqunu aqj

'J-

pue

'1-'Ju!w

Aq

pelorluor

sr unldas

oql

Jo

uotlelol aql

urnldas aqt

Jo

uoqelol

aql atelnbaS

saua!

u!u)

-eulqluof,JJ

e J^Pq

o] slueM

'JeAaMoq

'runrrJl

-)eq

JqJ

'runldas

Jr{l o1 JSOIJ sr aurosoruorq)

Jql

Jo

JruJnbJS

SnurruJal Jqt

uJqM elqelre^e

uorleurquoJar

JrJrJJds-Jlls

p

sr sJeql snqJ

'uorlJPJJ

eqt

uoddns louupJ

1l

'uorSJJ

slql

Jo

aprs

-lno

pe^ou

sl

1l

JI

1ql

0€-

Jo

uorSr.r

p

ur

pJreJol

Jq

tsnrrr

JJuenbJs

n8rcl

try

Jqt

'uorlrppp

uI

'uorsrlrp

IIer

pup

uorleSar8as

aruosotuoJqJ loJ

partnbar

sr

teql

runtdas

Jql ur

pJteJol

uratord e

'>ISt{

}o

aluasard

aqt ur Lpo s-rn))o

?aleMoq

'slupurqruo)ar

aa,r8 ol uorDunI

ar{l

Jo

uorlnlosJ5

'r{1lualsrsuor

raqloue

euo

purJ

ue) J)uanbJs

lJB

-Jet

Jqt

yo

satdor o,vrt

aql

,l,roq

sureldxJ

qJrqm

'aruanbasfp

eql rJAo

sassed >goJ uorlerrldar

eqt JetJe uoos

urJoJ leu xaldruoJ

eqJ

'ureql

uaJMlJq

uolnunILeprtloH

e ruJoJ

pue

saruanbas

Itp 1o

lred

p

ot

pulq

uer

)tJX

'It'ri

*sfi*Ij'

ur

pazrrpruurns

eJp sluJle

IUeAJIJJ

aq1

'de,t.

Sur

-lseJelur

ue ur uorsrlrp

IIJI

ol

palpler

sl uatsls

JaX Jqt

Jo

JSn aqJ

'JruosouorqJ

eqt

;o

uor8ar

snurrurJt

Jqt ur

pJtplol

sr

lpql

/p

pa11elatls

lJ8ret

dq-gf P uo

tf,e

q)lqm

'OrJX

pue

)rax

'sJSeuqruoJJJ

o1!1.l

Jo

slsrsuo)

srrtrl'llu

g, q

'ruJl

-s.,(s

uorleurqruoJJr

rryrrads-a1rs rJX

Jqt ssJSsod

sJurosoruorq)

relnJJrJ

qlrM

erJalJpq

lsow

'reurp

purseld

e se ,{.eat Jtups

Jqt ur

uorlnloseJ JAerqJp

ol

palnbar

sl uorteurqruoJaJ

puoJJs

e

'JSeJ

srqt

q

'JleJpdJs

louupJ

sJuros

-orrroJq)

ralq8nep

Jql

'luJAJ

uorleurqruoJJr

e

qJns

ureq

seq ereqt

JI

'JII^J

uorterlldar

e

.{q

parnpord

sauosourolqr

ralq8nep eqt uJJMteq

sJn)Jo

uollpurquo)Jr

snoSolouoq

Jr

'rJAa

-Moq

'peJnpord

aq

IIrM

rrrurp

V

'sllJt

ratq8nep

Jql 01 ale8ar8as

uef, seruosoruoJqJ

lJlq8npp

aleredas

Jql

pue

'rualqord

ou

sr eJJql srnJf,o

uorteurquoJJr

ou

JI

'uortB8ar8as

s1r

page

Laqt

Moq

sMoqs

r;'d.;

sHElsIj

leuosotuoJqJ

Ierrapeq

Jql

qlrM

rnJJo

uer

luJ^J

Jo

sadz{t aures

:q1

'uorlrpuoJ

JrrJurouoru

Jql serolsJJ

leql

uorleurqruo)JJ

Jpln)elourpltur

up Josuods ol

seJuJnbas relnrrued

uodn

pe

leqt

sruats.ds uo;l

-pulquroJer

rrynads-alls

J^eq uaqo sptuseld

'paJJJ

slql

DpJJlunoJ

oJ'lleJ

ralq8nep Juo

ruorJ

tsol

Jq

Ilqelrnaur

tsnru

JJolJJaql pnuseld

aqt

pue

'ateBarBJS

ot

lrun

auo

seq Lpo

1ar

Jqt

leql

supaur uorleurquorar

,{q

JJrutp

p

Jo

uorlpruJoJ

'paterlldar

lsnI

seq

leqt

pnuseld

Ldor-a13urs

e

Jo

eseJ aluJrlxe

eql

q

'slrun

Surle8ar8as .d11er

-rs,{.qd

Jo

JJqunu

Jql

setnper

tuJAJ

ue

qJnS

'surJoJ

JrrJrrrlllnru

raq8rq

aleraua8

uef, uorleurq

-ruoJJJ

rJqunJ lJIrrD

JuJrurp

p

salpJJuJB sJIJJT)

o1!{1 uas1!\lJq

lUJ^e

uorleurqruof,JJ

rplnJelou

-ra1ur

a13urs

y'saruanbasuo)

Jql selprlsuoruJp

*t"J.{ }'*i{l*g{

'aurquroral

o1 elqe are ,4aql

l1nsar

p

se

pue

'saruanbas

VNO

erups aql

Jo

lsrsuoJ

unrralJeq

e ur

puuseld

e;o

sardor aldrtlnur aq1

a1r^3

11a3

eql o]

papauuol

sI uoqelrldau

lPUepPS

'uoqeurquolar

rgnads-a1rs e

[q

srauouou

oMl

olut

pa^losa.l

aq uel srql'alnlalour luourrp e16uLs e saleraua6

'la^aMoq 'luala

uoqpurqulolal

pazrleleue6

V

'sllor

rel

-qbnep

o1 alebarbas

1eq1

srelqbnep

luaulouour oml ornp

-ord

ot salerrtda.l auosoruorqr lelnlrLr

V

x,{'fl.

IH{}. Ij

Offi

eleberbes

so ur osoruor rl c r e1q 6ne6

'illl*

.irxrr,

sorrjosouro.r r.lc

saseolo.l

uorleurquocoJ

crycads-e1rg

poureJlsuoc

sr

]uouro^ot\

ueoe

o^or.lr

sourosourorr.lc

ralqbne6

uorlEulquocarleraueg uolteurqulocoloN

\,/

\,,

\/

oruosoruorr]c Jelncrc euo sPq

llac

'slaur00rl0

urolj slrun

lenpnrpur

sosealal

uorleu rquoler relnlelouerlur

pue'srour

rp olu r srourouom

sablau uorleurquloler rplnlalouuolul

#I'l.i :HglSIj

alcjrc cuoulo

-

/I

U]IdVH] 9I'

LI'

sauosorrolqJ

aqlJo

uorlelpda5

sanlonul 6uruoL1L1e6

g'11

leql

serlr^rlJe ro

vNQ

01

purq

]pql

sural

-ord

apnlrur

plnor qJIqM

'uorle8ar8as

esner

ter{t

(s)uralord

Jqt loJ

Jpol

tpqt

saua8 ur rnJJo

plnoqs

suorlelnur Surt:e

-suau

:ssJJord

uorlrued Jql roJ

staSrel

eql

Jre

leql

saJuJnbas

VNe

ur Jn)Jo

plnoqs

suorlplnru

3ur1re-sl.r

:sJSSelf,

o,l.rl

pulJ

ol

Dadxa

JM'JleJ

JJe

Jlestr

ssarord

uottttred

Jql

IJaJJE teqt

suortetntr

l

o

's1ar

ralq8nep

luereJJrp

olur uaql ale8ar

-8as

ot rlqe

eq 01 JJpro ur ^dqerrSolodot

sJurosourorqr

s1r alSueluJsrp

ol JIqp

eq

lsnlu

unrJJlf,eq

eql

leql

sn sllJl srr{J

'seruosoruorqr

tatq8nep

qloq

Sururet

-uoJ

IIaJ

P

pue

IIeJ

elealJnue

ue saseJlJr

ueql

uorteuroJ untdas

'lla)-prur

tp

sspru

a8rel

a13urs e ur

peterol

sl

VNe

Jqt

tpql

tlnsJr

eql

qlplr

'Surle8ar8as

uor; seruos

-oruoJq)

ratq8nep

aql

luanard

suorl

-ptnu

Jqf

'rJqtoup

auo

qSno.rqt

spupJls

y|{q

ssed ot Itlllqp

Jqt

qlrm

saru.dzua

-sJspJJruosrodol

ro; Surpor saua8 ur

deu Suruortltred

SurtraJJe suorlelnru

')sll

Jo

eruesard

aq1 ur fi1uo

lr

anlosal uer

]nq

'alts

{!p

aqtte uoLl:unf r\epqlog

e so]earl .lax

'sauosouroiql

polutl

oMl saleerl

lua^a

uorlPurquotol

v

i;

I

.1

i.

!ijilli.l:l

lsow

'snurural

JQt

;o

dlruDn aqr uI

Jeqlo

qlpa punoJp palloJ

JJp

t€ql

suot8ar

VNCI

Jo

Sur18ueluastp sarnbar

sIqJ

'uoll

-eururJel

3umo1o;

ale8ar8as uer

[aqt

leq]

os JJqlouE Juo IUOJJ

paseelal

eq

lsnur

sJruosoruo;r-{)

.ratqSnep

o,r,rl JLlf r

:Suruortrued

radord ro;

parmbar

eJp

luala

Jo

saddl

o^,t41

'suuoJ

runldas eqt

qJrqm

te

uorlrsod aql

Jo

JpIS JJI{Ua

uo sJAlesureqt

pulJ

sJurosoruoJqr ratq8nep

o.trt eqt

qrlqm

Lq ssarord aql

sr Suruorllued

'suoLlsod,/E

pue

r

/l

eW ol lalual-ptur

aql uolj sluaueAoul

lonlqe

aleu Sauosouo.l{l

o

'auPlquau

lPu0llPq

rauur aql ol

paqleilp

aq r\eu surbuo

uorqdag

r

sauosoruoJql

aql

Jo

uor+Predas

sa^lo^ul 6uruoqrpe6

('saruosoruorq)

Jql

Jo

uoueredJs

seAIoAuI

3uruor111re4

'8'ZI

uoltJeS eeS)

'uoltelnJods

Surrnp arodsard

aql otq

tuaruuedruoJ

Jaqlou

eql ruorJ

ygq

sgodsuerr

gJIIodS

leqt,{e,u

arues

aql

ur

'runldas

aqt

q8norql

vN11

drund 01

prsn

Jq

plno)

srr4l'lritz'

/rl

vN61

3uo1e alerolsueJt ol

esn ueJ

tl

qJrqu Llrnrtre esedJv

ue seq osle

1I

'sJeruouour

o^\l

olur JaIuIp

e JAIOSaT 01 JaX esneJ

ot

sr Juo

'suorlJunJ

oAr1 sPq ulPluop

pulruJal-J

sU

'runtdas

eql ol

pJZIIpJol

Jq

ol

U

sJSneJ

pue

eueJqruJru

eql

qll1!\ pelPlJossP

sI

uleurop

IpurruJJl-N

sU'uralord

euerqruJlusuerl

a8tel

e sr

>JStd

'runldas

Jqt

ot

pelJJuuot sI uollnlo

-sJJ

teql

os

pJ^loAJ

seq ruals,{s

Jql

leql

slsaSSns

q)lqM

'esJal-JJIA

pue

Soloruoq

>IStd

ue JAPq

sLe.l,r1e tuats.,i.s

JJX aql J^eq

lpql

elJatJeg

'uats'(s

JeX eql ol

pasodxa

are,{.aqt

JreqM

'runldas

eql

Jo

,{ltutrt,r Jql

uI ulerueJ

e1

rusr{l sa)roJ

SII{J

'peluJo}

uJeq spq JaIuIp

p

Jr

pJureJlsuor

aq

rleur ?J^eMoq

laqloue euo

ruor;

uede

enoru ol

saruanbas

lueAJIeJ

aqr

Jo.dtl

-ilqp

eqJ

'Jeqloup

euo uorJ

lrede

a,rour sJruos

-oruoJqJ

oMl

eql

'uolleulqluoJeJ

ou uaeq

seq

eJaql

JI

'uorlerrldar

JJIJe uoos

suels sJruosour

-oJq)

Jo

uoue8ar8as

leqt

aq

r(eru:aMsue

JqJ

zrJrurp

e olul

paurq

-ruoJar

uJJq

Jneq ro

sJeurouour

]uapuadaput

se

tsrxJ

sJurosoruorqJ

ratq8nep

aql raqtJqM

n,tou>I

uratsui.s eql sJop

anog

(lrarutp

aql eleJrl

plnoM

uorteurquorar

rrynads-e1rs

eqt asIMJJqlO)

'JJrurp

e alBraua8

01

lue^e

uolleulquoJer

IeJJ

-ua8

e uaaq

,{pearle spq eraqt

uaq,tt

,{1uo uotl

uorlnlosoJ rol

pernoor

sr

ysll

eus uorleurqu_jocor

lP

polurl

are seLuosoruorqc

teqt

ssJ)oJd

lensnun

ue

sr srqJ

'(17'1

1

arn

-3r4

aas)

tuaulredurol

arodsaroJ

Ileurs

Jql

otut

pate8ar8as

aq

tsnru

JruosoruoJqr

ralq8nep

euo'stlqqns

snilDog ur

uorlelnJods Surrnq

'uorlPrlrul

JoJ

pJqtPtle

Jq

tsnru

ur8rro Jqt q)rq,vr

ot Juelqueu

aql

uo JrnlJnrls

e Jq

plno)

atrs qla,,ror8

JqI

'uollpJ

-ldar

;o

uorterlrul Jql qlr,lr

araJrJtur

teql

suorl

-e1nu

Aq

patra;;e

aq Leru

suorlJerl JuprqruJur

eseql ur

luasa.rd

suralord

ar{J'snururJl

Jql

pue

'>lJoJ

uorlpJrldar

aql

'ur8rro

Jql

rpau srJ>lJeur

rrlaua8 ur

pJrl)rJue

Jq ol

puel

qf,rqu

'suorl

-reJJ

JueJquau

ur

punoJ

eq upJ

vNo

IerJaDeg

'1)eJIpUr

SureurJJ

J)uJpr^a eql

lnq

'auerqruJur

eql

pue

vNC

IerJJllPq

uaaMleq slsrxJ >lurl

Ief,r

-s.{qd

e

teqt

sreaL:o;

patradsns

ueeq seq

tI

's1ar

ratq8nep

Jql

Jo

slJtuJJ

Jql euoleq

IIrM

leql

suorlrsod aql

le

sesseur

pasuapuoJ

ruJoJ 01

ruJql sasneJ

qJIqM

'c!Ig{nw

ol alels

pJsuepuoJun

up

ur

pessed

uJqt

pue

'sJserJluosrodol,{q

palSuetuJSrp

are'uor1ell1dar

uoJJ aBJJUJ sJurosoruorqr

ralq8nep JqJ'snteJ

-edde

uotlerrldar

aqt

q8norqt

ssed

ot JepJo ur

pJSuJpuoJJp

Jq

tsnru

1I

'pJuoursod,{11er1uar

sr

JruouJB

leluared

aqJ

'lJporu

luJrJn)

e sMor{s

I

i-1"1.

:iHtt*j:l'uorlesuepuoJ

qlrM pJl]euuof,

aq

Leru

ssarord

Jqt

lnq

'llJl

eql ur

pauorlrsod

are saruoua8 ,tvroq puelsJepun

lou

op

ruls

a^A

'uorle8ar8as

s,rorolle

snql

pup

uorlesuJpuof,

Jo

atels rado.rd aql

erotsat

ol sdlaq z{trsuap

gor

-radns

ur eseerJur

Surlpsar

Jql

lslroJrJdns

Jnrle

-3au

Surxelal

uoJJ sJSeJJruosrodol

Surluanard

.dq

patesuaduor

aq upr

lJeJJp

aq1

',{pa

-dord

ale8ar8as

o1 elnlreJ

Jo

asne)

Jqt sr uorl

-JunJ

srql ur

DeJap

v'Jurosourolqf,

Jql esuJpuoJ

ol usrupqJeru

Surlrorradns

p

sJsn

tI

'sursuJp

-uor

rtlo.drp>lnJ

ol snoSoleue

ursuepuoJ

e

Jq ol

pJJJplsuor

sr

xaldruor

{gg{nw

rqr

pue

g>lnwpup

f>lnw'suratordraqto

oMt

qtpu

xald

-uroJ

e suJoJ

g1nw

'I

JspJJtrrostodot

ro; sapor

teql

aua8 eqt'Vdll

ur suorlptnur

Lq

passarddns

Jq

uer

{4nw

ur

suorle}nur

Jtuos

tpql

,{ranorsrp

Jql spM

gInW

Jo

JIot

Jqt olur

tqBrsur

aq1

'prJelJeq

rJqto

ur

punoJ

uJJq osle

a^pq

sutalord aTI-)WS

'(sursuapuo3

Lq

pasne3

s1

uorlesuepuo)

Jurosourolq)

'g'I

€

uorllJS

aJs)

sJruosoruoJqr

rrlor(re>lna

raqlaSol

Surploq ur

pue

SursuapuoJ

ur

pJ^lolur

JJp

lpql

suralord

()ws)

sJruosourorqJ

Jo

aJupualureru

lptnlJnrls

yo

sdnor8

oml aqt

se uortezrueS,ro

Jo

ad,,(t

Iera

-ua8

arues aqt

seq

qrlqM

'urato-rd

re1nqo13

(q1

gg

1)

a8rel e roJ

srpor

g4nut

aua?

aql

'adola.t

ua

eql ol JruosoruoJqJ

aql Surqreue qtr,r,r pJ^lonur

aq

plnoJ

lrnpord

JSoqM

(2101)

uatotd

auerq

-ureru

JJlno

umou>l

e ro; aua8

Jqt

qtl,lr

lelnuJpr

sr

V4nu,t

aua8 aq1

'sJruosoruoJr{J

Jql saleBJJ

a1:43

11a3

aql ol

papauuo3

sI uo!]prqdax

leualreg

/I

UlldVH)

gll

'sproallnu

ralq6nep

aql sasuapuoler

1eq1

snleredde aq1

1o

luauoduor

1eL1

-uassa

up sr

€)nhl

'uoqerqdar

6uunp

pasuapuolop

souoloq

proallnu

leluered

a16uLs e

lo

VN0

eq1

{{.'/t 3#{l*Ij

-8as

leql

sntBredde Jqt

Jo

stuJuodruor,{;rtuapr

.{eru z{aql

pue

'lpqlJl

tou

Jre saua?

4nut

aql ur

suortetnw

'Surle8ar8as

Jo

pealsu

runtdas aql

JO

JprS aIuPS aql UO UrerUeJ SJUTOSOTUOTqJ

Jal

-q8nep

qlog

:zbuanbeJJ

paseeJJur

qrnru

e

1e

Lua

-3ord

alealrnup ot JSrJ a,rt8

qrrq,vr

'ssep4nw

erll

Io

suortptnru,{q

paldnrratur

sr uorle8al8ag

'suorleJol

rr;nads oJ sJruosoruorqr eql

anoru

.,i.eru

-stsaqlu,{s

uralord saJlnbar

leq1

auo-ssarord

alrDe up

teqt

stsaSSns srql'uoqe8uola

adola.tua

rJqunJ ,{.ue;o

aJuasqe Jql ur suorlrsod ralrenb

eql ol Jloru

seruosourorqr aqt'q8noqt'JurnseJ

ot

peMolle

sr srsaqtuLs uratord uJqM

'uor11sod

11e]-pru

Jqt ot JSop

urpural

pue

ale8ar8as 01

IreJ

sJruosouroJq)

aql'uoueJrlder

;o

uorleurruJel

Jql

eroJaq

pJtrqrqur

sr srsJqtu^ds uratord;1

'q18uel

11e]

eqt

Jo

sJJtJpnb aarql

pue

ragenb Juo

tp

suorl

-tsod

leur;

rraql 01 stueruJ,roru

ldnrqe

yo

alqeder

eJe seurosoruorqf,

palelldar

Jql

'eJouJJql

-Jnd

'aJpJrns

IIJ)

eloqM eql JeAo dlsnoauaSora

-laq,raor8

euerquJru

pue

IIeM IIe)

eql

'DeJ

uI

'uede

uraqt Surqsnd snql

'sauosouorqf,

o,r,rl

Jql

JO

Salrs

luauqJelle

Jql ueJMlaq

Ierrelelu

Jo

uoruJsur .{q

s.l,ror8 adolanua eqt

teql

palsaS

-3ns

g'41

arn8rg ur umoqs uorle8arSas

auos

-otuoJq)

JoJ

lJporu

Jql

Jo

ruJoJ

leur8r.ro

aq1

'sluetnur

uolllrred

01 relrurs sr

leqt

ad.{.touaqd

p

eleq eroJ

-Jraql

pue

'(sraruouolu

o,tvrl

Jo

ppJlsur

uortrl-red

01 JJrurp auo Lluo seq

11ar

8ur

-pI^Ip

Jql asneraq) ssol

purseld

JSpaJ)ur

sualsLs uorleurqtuoJa:

tryoads-alrs

pnu

-se1d

ur suorlelnu'uorlrppe

uI'Jruosotu

-oJq)

IerJJlJeq

aqt ur

punoJ

ueaq JApq

suorlrun;

Surl)e

-suglj

r{po

lnq'sprurseld

Sutuotltued

ro; alqrsuodsar sruals,{s

aql

ur

punoJ

uJJq J^pq

uortplnur

1o

sad,{t

qlog

'peq)ene

aq

tqSnu

vNo

rprqM 01

adolanua Jqt uo

suortpJol aql

IoJtuoJ

61,

ualsns 6uruor1L1e6

p

e^eH

splurspld fido3-a16ur5

6.11

'5;od

alrs

1abre1

aq1

pue gtod

pue

ynd

sauab

1o

slsrsuor

ualsfis uoLlebarbas

uourLuor

V tt"I; ]Sn$ru

Sred

gted yted

x--+-

rJol rJrql

Jql

Jo

^up

Jo

suorlalJ(

'yN(

uo alrs

Srud

rqt 01 spuq

q)q/!\

'grPd

ot spulq

1I

'rspdJv

up

sr

Vred

'seue8

omt

aql

Jo

rupJJtsuMop

tsnf

palerol (gtud)

tuauala

8ur1re-sD

e

pue

(gtud

pue

ynd)

oolSupe-suaq oml ere JJeql

z(1ertddtr

'rit"{l

IHfl}Ij

ur

pazrJpruruns

JJe ruJl

-s^s

uoruurol

e

Jo

sluauodruoJ aq1

'sptuseld

ur

pJrJuuapr

lsrr;

eJaM Euruorlrlred ur

pello^ur

suoltJunJ

'1reJ

uI

'11ar

ralq8nep

luerJJJrp

e o1

paleSar8as

aro;araql eJp

pue

'uorsrlrp

IIJJ

te

runldas er{l

Jo

sJprs

alrsoddo uo sJ^lJsueql

pur;

sardor

alerqdnp Jql

teqt

eJnsue o1 sruals,{s

Suruorlrued JJrnbJJ

spuuseld

.ddot-a13ur5

'erJalJpq

.{uaSord

Jo

JJqunu runrurxeul

aq1 ot

pruseld

aqt

yo

uorlpnlJdrJd arnsua

dlaq

ol

:asodrnd

aues aql

e^res

ile

to^JMoq

'uoll

-nlo^a

Jo

sruJal uI

'luJle

uortrtred aql Suueln

-3ar

qtr,r-,{ltJJJrp

peuJaJuo)

JJp sJeqto

sEaJJqM

^dprartput

tJp

sluels^s asJql

Jo

eruos

'lpnl^ins

sll

JJnsua

o1

z(lluapuadapur

8ur1re

1e

'sadL1

luaraJ

-Jrp

Jo

ueqo

'sruats^ds

IeJaAas

drrer

ol

pnuseld

e

roJ uoruruoJ

sl

U'uorlelndod

prrapeq

e ur

prru

-se1d

e

Jo IpArAJns

aql aJnsuJ 01

pesn

a.rp

ruslup

-qJeru

Jo

saddl

pranag

'JIJstr

uorteJrldar uodn

pe

lou

op

leqt tnq

/ssol

Jo

Lruanbar;

eqt

eseaJ)ul

lpqt

suorletnu dq

perJrluapr

eq ueJ uorle8ar

-8as

pnuseld

IoJluoJ

teqt

sruatsr(s aql

'(pruseld

^ddor-a13urs

p

JoJ uala

'1errdL1

sr uorsrnrp

IIJJ

rad

r-0I>)

ssol

Jo

selpJ Mol Lrarr

qlrm

suolteln

-dod

prrapeq

ur

pJurelurpur

aJp spruseld

'q-0

I>

;o

saouanbaJJ

le

spruseld

yo

ssol Jql ul

tlnsJJ

IIIM

sllal :alq8nep ot sprruseld;o uortnqrrlslp

Te)

-llsllels

e UJAJ JSnBJeq

'z(rpssarauun

sJuoJaq

rualsds

uorlBBarBas JArlJe ue'(runrralreq

rad

g1<

'arrtrerd

ur)

q8noua

lear8

sr reqrunu

eqt

JI

'tstxa

ol

sulSlro

yo

lood

e s,vt.olle

ler{1

rual

-s.4.s

uoue;lldar e aneq sptuseld.{dorr11nry

'sllrr

relqdnPp

tuereJJrp

aqt ot

pale8ar8as

are sur8rro ratq8nep

aql uJql

pue

'JJuo

paterrldar

eq upJ ut8rro

a18urs

y'euosoruorqJ

lerreDeq

aqt

Surura.no8

uollerqdar

;o; uralsr(s

aql

Jo

esoql 01 JEIIIUTs

are saruanbesuoJ

asoqm

IoJluoJ

uorlertldar

ro;

rua1s,{.s e a^er{ sprurseld

,{dor-a13ut5

'selnJJIoru

prurseld

Jo

raqunu

Jq1 se aIUES aql st sur8

-rJo

Jo

Jaqrunu

eql

llnsal

p

se

pue

'uorrldar

a1E

-urs

e

Jo

slsrsuof,

pruseld qreg

'runlJaDeq

aql ul

luasard

eq uet sur8rro

Lueru

lroq sJulruJJlap

uorlerqda,r

3ur1er11u1 ro;

alqrsuodsar uralsls aql

'tusru€qJaru

IoJluoJ

uolterqdar

1o

ad,{.1 eql

Jo

aruanbasuor

e,{.perurrd sr raqrunu

.,{.do3

'JIITOSOUTOTq)

Ierrel)pq

rad

(OZ

o1

91

,{1ertd^1) req

-runu

)usrJJllpJeqJ

p

ur

lsrxJ

spnuseld

z(dorrr1ny11

'unrrelJpq

Jad

pruseld

aqt;o

sardor

IeJeAes

aJp ereql

1pr{l lpsal

eql

qflzu'alrb

1ar

rad sluJ^a uoneutut

aldu

-lnru

Mollp srualsds

IoJluoJ

z(dorqp141 .

'JruosourorqJ

IPrraDeq

eql

qllM drlred sulelulPur

Lla,rrDa;;a

pguseld

.{.dot-a13urs

V'uols

-hrp

IIal

rad uorlerrldar

auo ul

lpser

pue

JruosoruoJqJ

lerJalJeq

eql

Jo

lEql

alqueseJ

srualsLs

IoJluoJ

.{dor-aputg o

:Jaqrunu

Ldoc

rrlsr:elrpJpql

e

lp lsoq

IeIrelJPq

sll

ur

peureturpru

sr

ptuseld yo

ad,{1

qreg

'ruatsds

uorlerrldar

yo

ad,{1 slt uodn

spuadap UoISIAIp

le

sllal ralq8nep

qloq

01

pelnqrrlslp

sI

U

leql

eJnsue ot

sJsn

pnuseld

e

lBql

rualsds

yo

ad.{1 aq1

'uorsr.^r.p

e r{q

parnpord

s11er ralqDnep

luala#lp

o1

palebel6es

ale

spLurseld

alerndnp

lpql

olnsua

suLalsrts uoll!ilPd

'slaul0u0tu

elPteuabat

01 u

orlPu rq ur olar .lPlnleloulPllu

t

alelapu

n

lsql

sualsfs uorlpurquolal

rgoads-a1rs

a^eq spluseld

'sreuqlnu

raqbtq

pue

srourlp

saleleua6 sptuseld

rplnrlD uaamlaq

uoqpu

rqulolel sno6o1ouro11

'utbuo

auosotuolqr

lPuapPq

lad

Ador

prurseld

l< le lstxo

spruseld

r{dorqlngrl

'

ut6uo

aurosoutolqr

lPuelrPq

rad fdor

prLuseld

auo

lp

lstxe

spLurseld

r{dor-e16ut5

ruals^s

Euruoqqred

e aAPH

sprrusPld

Ado3-al6u15

']uarulredruor

arodsaroy

aql o1

q8norql

y1qq

sdrund

lpql

uoltJunJ

uolteJolsueJl

P seq

dlqeqord

pue

urnldas eqt

1p

pelerol sr uralord

aIIIodS

aq1

'ssarord

slql roJ

partnbar

s'g111ods

'saua8

uoltelnrods

aqt

Jo

euo

'urnldas

lueJSPu

aql

ssoJJe eluosoluorqJ

er{l

Jo

Je}sueJl

SJAIOAUI

'YNUur

rql

Io

uorssJJdxJ

stue^aJd

1pql

vNu

JSuJSrlup

llplus

e sr Jtoprlue aql lurelord

Jlxol

p

JoJ

vNuu

Jql

sr

leql

rrllDl e

sPq

Iu

pruseld

eqJ

'surrlord

3ur

->lrolq

pue

JJIIDI

Jo

slsrsuo) prurspld

c

Jql

^q

pJIJ

-rJeds

euo

'sruJoJ

snorJp^ J{el

suJts^s esaqJ

'L1alrurlapur

pruseld

aql

ureter ot

pJuruJpuoJ

sr uorlelndod

eqt

pue

'arp

Llqelr,raur

prruseld

Jql esol

lPq]

PrrJlJPq

snql

'llJl

Jqt

Jo

qlpJp

aqt

sJSneJ

e)uetsqns

JJIIr{ Jqt uJqt

pup

's,,i.etap

eloprlue eqt

tsol

sr

pruseld

Jqt urq6

'pr^rl

goqs

.dlanrlelJJ

sr

tnq

uorlJe

rJIIr>l s>lJolq

teql

eJuelsqns e

Jo

slsrsuo)

Jloprlup aql

seJJaqM

'alQets

,{.lanrielar

sr

leqt

eJuelsqns

rJIIr{ e sr uos

-rod

aq1

'Jtopltue

ue

pue

uosrod e

qtoq

sJrnp

-ord

ptruseld

Jql

leql

i i

"

!.

:liiiil!

j:t

ur

pJlprlsnllr

aldnurrd eqt

Jreqs

qrlqM

Jo IIe

'prurseld

e

yo

,,pJJnJ,,

sl

tl

Jr

sarp

IIJJ

p

teqt

JJnsuJ

ol s^p,t,t

[p]J

-AJS

aJe araqJ

'pnuseld

aql

sureleJ

tr

se Suol se

.d1uo

anr,l.rns uel

pnuseld

e 8ur,{.rrer runrJJl)eq

p

teqt

arnsua

,,'Llateredas

Sueq

a.,r,r ro raqtaSol

Sueq a.lzr,,

tpqt

srseq

Jq1 uo aterado qtlqM

'srualsLs

uoIlJIppV

'uortrtred

radord eJnsue

leql

sprruseld

lenprrrrpur

ur

stuatsz{s

luapuad

-apur

'a1dr11nu

Jo

a)ualsrxJ

Jql Lq

pazrseqdrua

sr spnuseld errldar

ureS

s11ar rrlq8nep

IIe

teql

Surrnsua;o pnuseld

eqt

ot Jlueuodurr

aq1

'stJeratur

snteredde slql

qJIqM qtrM

sluJuodruor

re1n11ar

aqt L;uuapr

ol

rq

IIIM

srsLleue

aql

1o

a8ets

txeu

rql

'a1od

aqt

ol ur8uo Jr{l Surzrlelol

loJ

alqrsuodsar

sr snter

-edde

rr;oads

p

lpqt

lsaSSns

stlnsJr

asaql

'a1od

eqt

ol ur8rro aql

Surzrlerol

snqt

'ur8r-ro

Jqt ol

esolf, saJuJnbas

spurq

gred

pup

unrJaDpq

eqt

Jo

salod aql ot

ezlleJol

grpd

pue

yred'snjuilsan

fij

iaqlruo) u1

'sa1od

allsoddo

aqt

ot

pate8ar8as

are

sJruosouroJqJ

Jql

Irlun

Surrred

aluosouroJqJ

tll

lualentnba

sntets

e Sururelureru

'sur8rro

pazrs

-aqluzls

L1a'rau pup plo

qloq

spurq

1god5

leqt

alqtssod

sl

tI

'ulSlJo

aqt

;o

zi.lruru^

eqt ur Jruos

-ouorqJ

erp

Io

o/o1e-

rano pasradsrp

JJp

tpql

satdor

a1dr11nur

ur

tuasard

sr

leql

aluanbas e o1

spurq

lgodg

'araqt

ur8rro

aqt Surzllerol

JoJ Jlqrs

-uodser

aq,,{eru

pue

alod

Jqt

lp

sezrle)o1

16od5

's1ar

Suqelnrods

u1

'ftytt

arnSrg

aas) atodsatoy

Jql olul

aruosouroJqr

ralq8nep

auo

ale8ar8as

ol JJnlreJ

p

Jo

JSnp)eq

uorlelnrods

tua.tard

oo1

Jseqt ul suorletnw

'dlanrlradsar

'fOods

pue

!o5

palle)

Jre traql's1uqns'8,

uI

'prret)eq

IeJJ^es

ur

punoJ

Jre

gJed

pue

vred

o1

peleleJ

surelord

'runtdas

Jqt

Jo

qtMoJB

.{q

pate8ar8as

arp

tuaru

-vJeue

Jo

selrs

Jql uJqt

pue-JueJqrueu

aql

uo

'aldruexJ

roJ-etrs

lerrsLqd

eruos ot

VNC

Jqt sJqJeDe

]r

tpqt

sl

{tluqlssod

JUO

'{se1

srql

saqsrldruorre xalduror

Ienprlrpur

eqt

Jo

uorl

-eurroJ

eqt ,ltoq

1ad,r,rou1

tou

op e14'lJqtoup

apf3

11a3

aq] ot

papeuuol

sI uorlelqdaS

leuapeg

Juo ruoJJ

uede

ale8ar8as salnraloru

vNq

o^,rl.l

leqt

ernsuJ ot

:luJrJJJrp

sr xaldruor uolllupd

eqt

Jo

JIol JqI

'(auosetul

ue ur srnf,lo uorl

-€urquoJeu

ppquef

'6I'6I

uortJJS eas) aurq

-urorJJ

o1 uJql JIqeuJ o1 sJlnlJloru

VNCI

o.&u

spurq

uorlerSalur epquel aSeqd Surrnp

{HI

uo

sJlqruessp

teqr

xalduo)

VNq-urelord

aq1

'luarsueJl

lnq

IprluessJ

sr Jrnl)nJls xalduor uorl

-tued

aqt

qll^{

Vtpd

Jo

uortJeretur aq1

'L1anr1e

-radoor

purq

ol

gJed

Jo

sJerurp JJqunJ sJIqeuJ

sIqJ

'gred

Jo

rJrurp

p

qlrM

raqteSol

CHI

Io

rJrurpoJalJq e Lq

punoq

t

gnd

uar4,tt

palerlrur

sr

uorleuroJ xalduo3

'iii'

j.i.

:+!Jil*I*

ul

pJle)lpur

sp

'sJtrs

gxlq

pue

yxoq

paprcdas

eqt o1 z(1sno

-euellnlurs

purq

ueJ

gled

lpqt

os

yN(I

Jql

puJq

ot sr

CHI

Jo

JIoJ ar{J

'JleJJns

Jqt uo

padder,,n

sr

VN(I

qllqm

ur

arntJnlts

a8rel e uroJ

ol

u(tpeder

eqt seq

teql

JaurpoJJlJq

p

sl

JHI

'(eruosoru

-orqr

lsoq

rqt otur

VN(

epqruq a8eqd;o uorter8

-alur

Jqt ur

pJAIoAur

sl

teqt

eJnpnrts e SuIuroJ)

pJIaAoJSrp

lsJrl

se.&\

1l

qJrqM

ur elor Jql JoJ

pJrupu

sl

U'rol)p1

lsor{

uorle.rSalur Jrll sl

{HI

'gre4

u(q

punoq

are

leq]

gx1q

pue

vx)q

pJIIeJ

sJJuenbas Lq eprs rJqlra

uo

pJ>luplJ

sr

pup

Jlrs

Surpurq-gg1 Jql SurureluoJ e)uJnbes

dq-tg

e sr

gtud'xaldruor

uorlrlred eqt

peller

sr

gwd

qtlM

{HI

pue

grpd

;o

xaldruor

eqJ

'Jrnl

-JnJts

Jql

;o

ged

uJoJ ol Jlrs srqt

te

spurq oslp

'9111 'uralord

IerrJDeq

V

's1ar

ratq8nep

atrsoddo

ot sardor

pruseld

aqt sale8ar8Js

leql

Jrnl)nns

p

sateJJ)

t1

ot uratord

gJed

eqt

yo

Surpurg

'1ar

)Ilo^Je>lnJ e ur eJJuroJlueJ Jql o1

luale,rrnba

sr

tpqt

prurseld

eqt roJ eloJ e s[,e1d

gtad

'selrs

tsurlf,P-s,

Io

sauaS

Surpuodsarror Jql uJJM]Jq sarSoloruoq

aluanbas

luel;ru8rs

ou JJp alaql

'sJrlrrelurs

11e

-rJAo

llJqt;o

atrds

q

'IU

pup

'Id

{

spruspld eql

JoJ

pazrrJlJeJpqJ

ueJq a,req ad.{t slql

Jo

surJl

-si(g

'pruseld

aqt

to

uorlrued radord

tua,r.ard

'purq

slaurp

Sled

ar0ur ueql

pue

'grPd

J0

rourrp

P

pup

IHI

Jo

lorurpo.la

-1aq

e

fiq

palprlrur

sL xelduLor eqt

'eprs

raqlra

uo salrs o]

purq

uer

Eled

lpql

os

lr

spuaq

pue gtod

le

VN6

ot spurq

IHI

uaqM

peuloJ

sr

xaldLuor

uorlrled aql

ii..

j.r

tSIir{:.:'

/r

ulldvHl

oz,

Lewin Benjamin (ed.) Genes IX

Подождите немного. Документ загружается.