Principles Of Development 5th Edition Lewis Wolpert Free Pdf

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
1
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
Ch«»i».
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
I
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
9
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