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Principles Of Development
Article January 1998
Source: OAI

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Principles of
Development
Lewis Wolpert
Thomas Jessell
Peter Lawrence
Elliot Meyerowitz
Elizabeth Robertson
Jim Smith

OXFORD
UNIVERSITY PRESS

Summary of contents
Contents
Text acknowledgements
Figure acknowledgements

XI

xviii
xix

Chapter' 1

History and basic concepts

Chapter 2

Development of the Drosophila body plan

31

Chapter 3

Patterning the vertebrate body plan 1:

axes and germ layers

89

Patterning the vertebrate body plan II:

the somites and early nervous system

149

Development of nematodes, sea urchins,

ascidians, and slime molds

185

Chj.

Plant development

225

Ch>,

Morphogenesis: change in form in

the early embryo

257

Cell differentiation and stem cells

297

Chi.

Organogenesis

339

Chrt-,-

Development of the nervous system

387

Ch.K. ,

Germ cells, fertilization, and sex

421

Cl-

Growth and post-embryonic development

451

CUt-

Regeneration

475

Cf

Evolution and development

497

Ow/

Glossary

525

Index

537

Contents
Ch .

History and basic concepts

Development of
the Drosophila body plan

The origins of developmental biology

Drosophila life cycle and overall development

2.1 The early Drosophila embryo is a multinucleate
syncytium

1.1 Aristotle first defined the problem of epigenesis

and preformation
Box 1A Basic stages of Xenopus laevis development

2.2 Cellularization is followed by gastrulation,

segmentation, and the formation of the larval nervous
system

1.2 Cell theory changed the conception of embryonic

development and heredity
1.3 Two main types of development were originally proposed

2.3 After hatching the Drosophila larva develops

through several larval stages, pupates, and then undergoes
morphogenesis to become an adult

1.4 The discovery of induction showed that one group

of cells could determine the development of neighboring cells
i.s The study of development was stimulated by
the coming together of genetics and development
1.6 Development is studied mainly through a selection
of model organisms

32
33
33

34

2.4 Many developmental mutations have been

identified in Drosophila through induced mutation
and large-scale genetic screening

35

Box 2A Mutagenesis and genetic screening strategy

for identifying developmental mutants in Drosophila

36

1.7 The first developmental genes were identified as

spontaneous mutations

11

Setting up the body axes

37

A conceptual tool kit

13
13

2.5 The body axes are set up while the Drosophila

embryo is still a syncytium

37

15

2.6 Maternal factors set up the body axes and direct

the early stage of Drosophila development

38

Box 1B Germ layers

1.9 Cell behavior provides the link between gene
action and developmental processes

16

2.7 Three classes of maternal genes specify

the antero-posterior axis

40

1.10 Genes control cell behavior by specifying which

proteins are made

17

2.8 The bicoid gene provides an antero-posterior

gradient of morphogen

41

1.11 The expression of developmental genes is under

the control of complex control regions

18

2.9 The posterior pattern is controlled by the gradients

of Nanos and Caudal proteins

42

1.12 Development is progressive and the fate of cells

becomes determined at different times

19

43

1.13 Inductive interactions can make cells different

from each other

22

2.10 The anterior and posterior extremities of

the embryo are specified by cell-surface receptor
activation

23

2.11 The dorso-ventral polarity of the embryo is

specified by localization of maternal proteins in
the egg vitelline envelope

44

1.14 The response to inductive signals depends on

the state of the cell
i.is Patterning can involve the interpretation
of positional information

23

2.12 Positional information along the dorso-ventral

axis is provided by the Dorsal protein

45

1.16 Lateral inhibition can generate spacing patterns

25

46

1.17 Localization of cytoplasmic determinants and

asymmetric cell division can make cells different
from each other

25

Box 2B The Toll signaling pathway: a multifunctional

pathway
Localization of maternal determinants during
oogenesis

47

1.18 The embryo contains a generative rather than

a descriptive program

26

48

1.19 The reliability of development is achieved

by a variety of means

27

2.13 The antero-posterior axis of the Drosophila egg

is specified by signals from the preceding egg chamber
and by interactions of the oocyte with follicle cells

27

2.14 The dorso-ventral axis of the egg is specified by

movement of the oocyte nucleus followed by siqnalinq
between oocyte and follicle cells

51

1.20 The complexity of embryonic development

is due to the complexity of cells themselves

1.8 Development involves cell division, the emergence

of pattern, change in form, cell differentiation, and growth

XII

CONTENTS

Patterning the early embryo

52

Box3A Polar bodies

93

2.15 The expression ofzygotic genes along

52

3.2 The zebrafish embryo develops around a large,

undivided yolk

96

55

3.3 The early chicken embryo develops as a flat disc

of cells overlying a massive yolk

98

57

Box3B Large-scale mutagenesis in zebrafish

99
104

57

3.4 Early development in the mouse involves

the allocation of cells to form the placenta and
extra-embryonic membranes

58

Setting up the body axes

108

Box 2C P-element-mediated transformation

59

3.5 The animal-vegetal axis is maternally

determined in Xenopus and zebrafish

109

Activation of the pair-rule genes and

the establishment of parasegments

61

110

2.20 Parasegments are delimited by expression of

pair-rule genes in a periodic pattern

61

3.6 Localized stabilization of the transcriptional

regulator (3-catenin specifies the future dorsal side
and the location of the main embryonic organizer
in Xenopus and zebrafish

2.21 Gap-gene activity positions stripes of pair-rule

gene expression

62

Box3C Intercellular signals in development

111

the dorso-ventral axis is controlled by Dorsal protein

2.16 The Decapentaplegic protein acts as a morphogen

to pattern the dorsal region

2.17 The antero-posterior axis is divided up into broad

regions by gap-gene expression

2.18 Bicoid protein provides a positional signal for

the anterior expression of hunchback

2.19 The gradient in Hunchback protein activates

and represses other gap genes

Box 3D In situ detection of gene expression

112

3.7 Signaling centers develop on the dorsal side

of Xenopus and zebrafish

115

Segmentation genes and compartments

65

2.22 Expression of the engrailed gene delimits

a cell-lineage boundary and defines a compartment

65

3.8 The antero-posterior and dorso-ventral axes

of the chick blastoderm are related to the primitive streak

117

Box 2D Genetic mosaics and mitotic recombination

68
70

3.9 The axes of the mouse embryo are not recognizable

early in development

119

2.23 Segmentation genes stabilize parasegment

3.10 The bilateral symmetry of the early embryo is broken

121

boundaries and set up a focus of signaling at

the boundary that patterns the segment
2.24 Insect epidermal cells become individually polarized

to produce left-right asymmetry of internal organs

73

in an antero-posterior direction in the plane of the epithelium

Box 2E Planar cell polarity

75

2.25 Some insects use different mechanisms for

76

patterning the body plan

The origin and specification of the germ layers

12 5

3.11 A fate map of the amphibian blastula is

constructed by following the fate of labeled cells

125

3.12 The fate maps of vertebrates are variations

127

on a basic plan
Specification of segment identity
2.26 Segment identity in Drosophila is specified by
genes of the Antennapedia and bithorax complexes
2.27 Homeotic selector genes of the bithorax complex

78
78
79

are responsible for diversification of the posterior

segments
80

2.29 The order of Hox gene expression corresponds to

the order of genes along the chromosome

81

Box 2F Targeted gene expression and misexpression

screening

128

have their fates determined and regulation is possible

Box 3E Producing developmental mutations in mice

130

3.14 In Xenopus the endoderm and ectoderm are

130

specified by maternal factors, but the mesoderm is

induced from ectoderm by signals from the vegetal region

2.28 The Antennapedia complex controls specification

of anterior regions

2.30 The Drosophila head region is specified by genes

other than the Hox genes

3.13 Cells of early vertebrate embryos do not yet

3.15 Mesoderm induction occurs during a limited

132

period in the blastula stage

81
82

3.16 Zygotic gene expression is turned on at

the mid-blastula transition

133

3.17 Mesoderm-inducing and patterning signals are

134

produced by the vegetal region, the organizer,

and the ventral mesoderm
3.18 Members of the TGF-p family have been identified

136

as mesoderm inducers

Patterning the vertebrate

body plan I: axes andgerm layers
Vertebrate life cycles and outlines of development
3.1 The frog Xenopus laevis is the model amphibian
for developmental studies

3.19 The dorso-ventral patterning of the mesoderm

137

involves the antagonistic actions of dorsalizing and

ventralizing factors
90
92

3.20 Mesoderm induction and patterning in the chick

and mouse occurs during primitive-streak formation

139

3.21 Gradients in signaling proteins and threshold

responses could pattern the mesoderm

140

CONTENTS

Chapter 4: Patterning the vertebrate

body plan II: the somites and early
nervous system

xiii

5.6 Vulval development is initiated by the induction

of a small number of cells by short-range signals
from a single inducing cell

199

Echinoderms

202

Somite formation and antero-posterior patterning

151
151

5.7 The sea urchin embryo develops into

a free-swimming larva

202

4.1 Somites are formed in a well defined order along

the antero-posterior axis

155

5.8 The sea urchin egg is polarized along

the animal-vegetal axis

203

4.2 Identity of somites along the antero-posterior axis

is specified by Hox gene expression

156

5.9 The oral-aboral axis in sea urchins is related

to the plane of the first cleavage

205

Box 4A The Hox genes

Box4B Gene targeting: insertional mutagenesis
and gene knock-out

158

5.10 The sea urchin fate map is finely specified,

yet considerable regulation is possible

206

4.3 Deletion or overexpression of Hox genes causes

changes in axial patterning

162

5.11 The vegetal region of the sea urchin embryo

acts as an organizer

207

4.4 Hox gene activation is related to a timing

mechanism

163

5.12 The sea urchin vegetal region is specified

208

4.5 The fate of somite cells is determined by signals

from the adjacent tissues

164

The role of the organizer and neural induction

166

Ascidians

212

4.6 The inductive capacity of the organizer changes

during gastrulation

167

5.14 In ascidians, muscle is specified by localized

214

4.7 The neural plate is induced in the ectoderm

169

by nuclear accumulation of p-catenin

5.13 The genetic control of e n d o m e s o d e r m

210

specification is known in considerable detail

cytoplasmic factors
5.15 Mesenchyme and notochord development in

215

ascidians require signals from the endoderm

4.8 The nervous system can be patterned by signals

from the mesoderm

173

Cellular slime molds

217

4.9 There is an organizer at the midbrain-hindbrain

boundary

175

s.16 Patterning of the slime mold slug involves cell

sorting and positional signaling

218

4.10 The hindbrain is segmented into rhombomeres

175

5.17 Chemical signals direct cell differentiation

in the slime mold

219

by boundaries of cell-lineage restriction

4.11 Neural crest cells arise from the borders
of the neural plate

177

4.12 Hox genes provide positional information

178

in the developing hindbrain

4.13 The e m b r y o is patterned by t h e neurula stage

179

Plant development
6.1 The model plant Arabidopsis thaliana has a short
life cycle and a small diploid genome

226

into organ-forming regions that can still regulate

Embryonic development

228

*
Development of nematodes,
sea urchins, ascidians, and slime molds

6.2 Plant embryos develop through several

distinct stages

228

Box 6A Angiosperm embryogenesis

229

6.3 Gradients of the signal molecule auxin establish

the embryonic apical-basal axis

231

6.4 Plant somatic cells can give rise to embryos

and seedlings

232

Box 6B Transgenic plants

234

:i

Nematodes

186

Box 5A Gene silencing by RNA interference

189

s.i The antero-posterior axis in C. elegans is

determined by asymmetric cell division

190

5.2 The dorso-ventral axis in C. elegans is

determined by cell-cell interactions

191

Meristems

234

5.3 Both asymmetric divisions and cell-cell interactions

specify cell fate in the early nematode embryo

193

6.5 A meristem contains a small central zone

of self-renewing stem cells

235

5.4 A small cluster of Hox genes specifies cell fate

along the antero-posterior axis

195

6.6 The size of the stem-cell area in the meristem is kept

constant by a feedback loop to the organizing center

236

5.5 The timing of events in nematode development

is under genetic control that involves microRNAs

196

6.7 The fate of cells from different meristem layers

can be changed by changing their position

237

Box 5B Gene silencing by microRNAs

197

6.8 A fate map for the embryonic shoot meristem

can be deduced using clonal analysis

238

XIV

CONTENTS

6.9 Meristem development is dependent on signals

from other parts of the plant

240

6.10 Gene activity patterns the proximo-distal and

adaxial-abaxial axes of leaves developing from the
shoot meristem

240

6.11 The regular arrangement of leaves on a stem

and trichomes on leaves is generated by competition
and lateral inhibition

242

6.12 Root tissues are produced from Arabidopsis

root apical meristems by a highly stereotyped pattern
of cell divisions

243

7.11 Vertebrate gastrulation involves several different

276

types of tissue movement

7.12 Convergent extension and epiboly are

280

due to different types of cell intercalation

Neural-tube formation

283

7.13 Neural-tube formation is driven by changes

284

in cell shape and cell migration

Cell migration

286

7.14 Neural crest migration is controlled by

286

environmental cues and adhesive differences

Flower development and control of flowering

246

6.13 Homeotic genes control organ identity

in the flower

246

Box 6C The basic model for the patterning of

the Arabidopsis flower

249

6.14 The Antirrhinum flower is patterned dorso-ventrally

250

as well as radially
6.15 The internal meristem layer can specify floral

251

meristem patterning
6.16 The transition of a shoot meristem to a floral
meristem is under environmental and genetic control

251

7.15 Slime mold aggregation involves chemotaxis

288

and signal propagation

Directed dilation

290

7.16 Later extension and stiffening of the notochord

occurs by directed dilation

290

7.17 Circumferential contraction of hypodermal cells

291

elongates the nematode embryo

7.18 The direction of cell enlargement can determine
the form of a plant leaf

292

Cell differentiation
and stem cells

Morphogenesis: change
in form in the early embryo

Box 8A DNA microarrays for studying gene expression

299

Cell adhesion

258

The control of gene expression

301

Box 7A Cell-adhesion molecules and cell junctions

259
260

8.1 Control of transcription involves both general

and tissue-specific transcriptional regulators

301

7.1 Sorting out of dissociated cells demonstrates

differences in cell adhesiveness in different tissues

8.2 External signals can activate genes

303

7.2 Cadherins can provide adhesive specificity

261
262

7.3 The asters of the mitotic apparatus determine

the plane of cleavage at cell division

263

8.3 Maintenance and inheritance of patterns of gene

activity may depend on chemical and structural
modifications to chromatin as well as on regulatory
proteins

305

Cleavage and formation of the blastula

7.4 Cells become polarized in early mouse and

sea urchin blastulas

264

7.5 Ion transport is involved in fluid accumulation

in the frog blastocoel

Models of cell differentiation

309
310

266

8.4 All blood cells are derived from multipotent

stem cells

312

7.6 Internal cavities can be created by cell death

267

8.5 Colony-stimulating factors and intrinsic changes

control differentiation of the hematopoietic lineages

Castrulation movements

269

314

7.7 Gastrulation in the sea urchin involves cell

migration and invagination

269

8.6 Developmentally regulated globin gene expression

is controlled by regulatory sequences far distant from
the coding regions

270

8.7 Differentiation of cells that make antibodies involves

irreversible DNA rearrangement

316

Box 7B Change in cell shape and cell movement

7.8 Mesoderm invagination in Drosophila is due to
changes in cell shape that are controlled by genes
that pattern the dorso-ventral axis

273

8.8 The epithelia of adult mammalian skin and gut

are continually replaced by derivatives of stem cells

318

275

8.9 A family of genes can activate muscle-specific

transcription

319

7.9 Germ-band extension in Drosophila involves

myosin-dependent intercalation

276

8.10 The differentiation of muscle cells involves

withdrawal from the cell cycle, but is reversible

320

7.10 Dorsal closure in Drosophila and ventral closure

in C. elegans are brought about by the action of
filopodia

8.11 Skeletal muscle and neural cells can be renewed

from stem cells in adults

321

CONTENTS

XV

8.12 Embryonic neural crest cells differentiate into

a great variety of different cell types

322

9.16 Drosophila wing epidermal cells show planar

cell polarity

363

8.13 Programmed cell death is under genetic control

324

9.17 The leg disc is patterned in a similar manner

to the wing disc, except for the proximo-distal axis

363

The plasticity of gene expression

327
327

9.18 Butterfly wing markings are organized by

additional positional fields

364

8.14 Nuclei of differentiated cells can support

development

329

9.19 The segmental identity of imaginal discs is

determined by the homeotic selector genes

366

8.15 Patterns of gene activity in differentiated cells

can be changed by cell fusion

330

9.20 Patterning of the Drosophila eye involves

cell-cell interactions

367

8.16 The differentiated state of a cell can change by

transdifferentiation
8.17 Embryonic stem cells can proliferate and
differentiate into many cell types in culture

332

9.21 Activation of the gene eyeless can initiate

eye development

8.18 Stem cells could be a key to regenerative

medicine

332

Internal organs: blood vessels, lungs, kidneys,

heart, and teeth

371

9.22 The vascular system develops by vasculogenesis

followed by angiogenesis

372

9.23 The tracheae of Drosophila and the lungs of

vertebrates branch using similar mechanisms

374

9.24 The development of kidney tubules involves

reciprocal induction by the ureteric bud and surrounding
mesenchyme

375

Organogenesis

369

The vertebrate limb

340

9.1 The vertebrate limb develops from a limb bud

340

9.2 Patterning of the limb involves positional information

341

9.3 Genes expressed in the lateral plate mesoderm

are involved in specifying the position and type
of limb

341

9.25 The development of the vertebrate heart

involves specification of a mesodermal tube that is
patterned along its long axis

377

9.4 The apical ectodermal ridge is required for limb

outgrowth

343

9.26 A homeobox gene code specifies tooth identity

379

9.5 The polarizing region specifies position along

the limb's antero-posterior axis

345

Box9A Positional information and morphogen

gradients

347

Development of
the nervous system

9.6 Position along the proximo-distal axis may be

specified by a timing mechanism

349

Specification of cell identity in the nervous system

388

10.1 Neurons in Drosophila arise from proneural clusters

388
390

9.7 The dorso-ventral axis is controlled by the ectoderm

350

9.8 Different interpretations of the same positional

signals give different limbs

351

10.2 Asymmetric cell divisions and timed changes

in gene expression are involved in the development
of the Drosophila nervous system

9.9 Homeobox genes also provide positional values

for limb patterning

10.3 The neuroblasts of the sensory organs of adult

Drosophila are already specified in the imaginal discs

392

351

9.10 Self-organization maybe involved in

the development of the limb bud

10.4 The vertebrate nervous system is derived from

the neural plate

392

353

9.11 Limb muscle is patterned by the connective

tissue

10.5 Specification of vertebrate neuronal precursors

involves lateral inhibition

393

354

Box 9B Reaction-diffusion mechanisms

10.6 Neurons are formed in the proliferative zone

of the neural tube and migrate outwards

394

355

9.12 The initial development of cartilage, muscles,

and tendons is autonomous

356

396

9.13 Joint formation involves secreted signals

and mechanical stimuli

357

10.7 The pattern of differentiation of cells along

the dorso-ventral axis of the spinal cord depends
on ventral and dorsal signals

9.14 Separation of the digits is the result

of programmed cell death

357

Neuronal migration

401

10.8 The growth cone controls the path taken

by the growing axon

402

Insect wings, legs, and eyes

358
359

10.9 Motor neurons from the spinal cord make

muscle-specific connections

403

9.15 Positional signals from the antero-posterior

and dorso-ventral compartment boundaries pattern
the wing imaginal disc

10.10 Axons crossing the midline are both

attracted and repelled

405

XVI

CONTENTS

10.11 Neurons from the retina make ordered

connections on the tectum to form a retino-tectal map

406

Synapse formation and refinement

409

10.12 Synapse formation involves reciprocal

411

interactions

Chapter 12: Growth and post-embryonic

development
Growth

451

12.1 Tissues can grow by cell proliferation,

cell enlargement, or accretion

452

10.13 Many motor neurons die during normal development

412

10.14 Neuronal cell death and survival involve both

413

12.2 Cell proliferation can be controlled by

an intrinsic program

452

414

12.3 Organ size can be controlled by external

signals and intrinsic growth programs

454

12.4 Organ size may be determined by absolute

dimension rather than cell number

455

12.5 Growth can be dependent on growth hormones

457

12.6 Growth of the long bones occurs in the growth

plates

458

intrinsic and extrinsic factors

10.15 The map from eye to brain is refined by neural
activity

Chapter ] 1: Germ cells, fertilization,

and sex
The development of germ cells

422

11.1 Germ-cell fate can be specified by a distinct

germplasm in the egg

422

12.7 Growth of vertebrate striated muscle is dependent

on tension

460

11.2 Pole plasm becomes localized at the posterior

end of the Drosophila egg

425

12.8 Cancer can result from mutations in genes

that control cell multiplication and differentiation

461

11.3 Germ cells migrate from their site of origin

to the gonad

425

12.9 Hormones control many features of plant growth

463

Molting and metamorphosis

465

11.4 Germ-cell differentiation involves a reduction

in chromosome number

426

12.10 Arthropods have to molt in order to grow

465

n.5 Oocyte development can involve gene

amplification and contributions from other cells

427

12.11 Metamorphosis is under environmental

and hormonal control

466

n.6 Some genes controlling embryonic

growth are imprinted

427

Aging and senescence

469

12.12 Genes can alter the timing of senescence

470

Fertilization

432

12.13 Cultured mammalian cells undergo cell

471

11.7 Fertilization involves cell-surface interactions

between egg and sperm

432

n.8 Changes in the egg membrane at fertilization

block polyspermy

434

11.9 A calcium wave initiated at fertilization results

in egg activation

435

Determination of the sexual phenotype

437

11.10 The primary sex-determining gene in mammals

is on the Y chromosome

senescence

13 Regeneration
Limb and organ regeneration

476

13.1 Vertebrate limb regeneration involves cell

dedifferentiation and growth

477

13.2 The limb blastema gives rise to structures with

positional values distal to the site of amputation

480

437
438

13.3 Retinoic acid can change proximo-distal

positional values in regenerating limbs

482

11.11 Mammalian sexual phenotype is regulated

by gonadal hormones

439

13.4 Insect limbs intercalate positional values

by both proximo-distal and circumferential growth

483

11.12 The primary sex-determining signal in Drosophila

is the number of X chromosomes, and is cell autonomous

441

13.5 Heart regeneration in the zebrafish does not

involve dedifferentiation

486

n.13 Somatic sexual development in Caenorhabditis

is determined by the number of X chromosomes

442

13.6 The mammalian peripheral nervous system

can regenerate

486

n.14 Most flowering plants are hermaphrodites,

but some produce unisexual flowers
ii.is Germ-cell sex determination can depend both
on cell signals and genetic constitution

443

Regeneration in Hydra

488
488

n.16 Various strategies are used for dosage

compensation of X-linked genes

444

13.7 Hydra grows continuously but regeneration

does not require growth
13.8 The head region of Hydra acts both as an
organizing region and as an inhibitor of inappropriate
head formation

489

CONTENTS

13.9 Head regeneration in Hydra can be accounted

490

14.6 Changes in Hox genes have generated the

elaboration of vertebrate and arthropod body plans

510

492

14.7 The position and number of paired appendages in

insects is dependent on Hox gene expression

513

14.8 The basic body plan of arthropods and vertebrates

is similar, but the dorso-ventral axis is inverted

514

14.9 Evolution of spatial pattern may be based on just

a few genes

516

Changes in the timing of developmental processes

517

14.10 Changes in growth can alter the shapes

of organisms

517

14.11 The timing of developmental events has

changed during evolution

517

The evolution of development

520

14.12 How multicellular organisms evolved from

520

for in terms of two gradients

13.10 Genes controlling regeneration in Hydra are

similar to those expressed in animal embryos

Chapter 14 Evolution and development

14.1 The evolution o f life histories has implications

xvii

500

for development
The evolutionary modification of embryonic

501

development
14.2 Embryonic structures have acquired new functions
during evolution

502

14.3 Limbs evolved from fins

504

14.4 Vertebrate and insect wings make use of

evolutionarily conserved developmental mechanisms

508

14.5 Hox gene complexes have evolved through gene

duplication

508

single-celled ancestors is still highly speculative