Chapter 47 Animal Development

Overview: A Body-Building Plan It is difficult to imagine that each of us began life as a single cell called a zygote A human embryo at about 6–8 weeks after conception shows development of distinctive features

Fig. 47-1 Figure 47.1 How did this complex embryo form from a single cell? 1 mm

The question of how a zygote becomes an animal has been asked for centuries As recently as the 18th century, the prevailing theory was called preformation Preformation is the idea that the egg or sperm contains a miniature infant, or “homunculus,” which becomes larger during development

Fig. 47-2 Figure 47.2 A “homunculus” inside the head of a human sperm

Development is determined by the zygote’s genome and molecules in the egg called cytoplasmic determinants Cell differentiation is the specialization of cells in structure and function Morphogenesis is the process by which an animal takes shape

Model organisms are species that are representative of a larger group and easily studied, for example, Drosophila and Caenorhabditis elegans Classic embryological studies have focused on the sea urchin, frog, chick, and the nematode C. elegans

Concept 47.1: After fertilization, embryonic development proceeds through cleavage, gastrulation, and organogenesis Important events regulating development occur during fertilization and the three stages that build the animal’s body Cleavage: cell division creates a hollow ball of cells called a blastula Gastrulation: cells are rearranged into a three-layered gastrula Organogenesis: the three layers interact and move to give rise to organs

Fertilization Fertilization brings the haploid nuclei of sperm and egg together, forming a diploid zygote The sperm’s contact with the egg’s surface initiates metabolic reactions in the egg that trigger the onset of embryonic development

The Acrosomal Reaction The acrosomal reaction is triggered when the sperm meets the egg The acrosome at the tip of the sperm releases hydrolytic enzymes that digest material surrounding the egg

Fig. 47-3-1 Basal body (centriole) Sperm head Figure 47.3 The acrosomal and cortical reactions during sea urchin fertilization For the Cell Biology Video Cortical Granule Fusion Following Egg Fertilization, go to Animation and Video Files. Acrosome Jelly coat Vitelline layer Sperm-binding receptors Egg plasma membrane

Fig. 47-3-2 Basal body (centriole) Sperm head Figure 47.3 The acrosomal and cortical reactions during sea urchin fertilization Acrosome Hydrolytic enzymes Jelly coat Vitelline layer Sperm-binding receptors Egg plasma membrane

Fig. 47-3-3 Sperm nucleus Acrosomal process Basal body (centriole) Actin filament Sperm head Figure 47.3 The acrosomal and cortical reactions during sea urchin fertilization Acrosome Hydrolytic enzymes Jelly coat Vitelline layer Sperm-binding receptors Egg plasma membrane

Fig. 47-3-4 Sperm plasma membrane Sperm nucleus Acrosomal process Basal body (centriole) Actin filament Sperm head Fused plasma membranes Figure 47.3 The acrosomal and cortical reactions during sea urchin fertilization Acrosome Hydrolytic enzymes Jelly coat Vitelline layer Sperm-binding receptors Egg plasma membrane

Fig. 47-3-5 Sperm plasma membrane Sperm nucleus Fertilization envelope Acrosomal process Basal body (centriole) Actin filament Sperm head Cortical granule Fused plasma membranes Figure 47.3 The acrosomal and cortical reactions during sea urchin fertilization Perivitelline space Acrosome Hydrolytic enzymes Jelly coat Vitelline layer Sperm-binding receptors Egg plasma membrane EGG CYTOPLASM

Gamete contact and/or fusion depolarizes the egg cell membrane and sets up a fast block to polyspermy

The Cortical Reaction Fusion of egg and sperm also initiates the cortical reaction This reaction induces a rise in Ca2+ that stimulates cortical granules to release their contents outside the egg These changes cause formation of a fertilization envelope that functions as a slow block to polyspermy

Fig. 47-4 EXPERIMENT 10 sec after fertilization 25 sec 35 sec 1 min 500 µm RESULTS 1 sec before fertilization 10 sec after fertilization 20 sec 30 sec 500 µm Figure 47.4 Is the distribution of Ca2+ in an egg correlated with formation of the fertilization envelope? For the Cell Biology Video Calcium Release Following Egg Fertilization, go to Animation and Video Files. For the Cell Biology Video Calcium Wave Propagation in Fish Eggs, go to Animation and Video Files. CONCLUSION Point of sperm nucleus entry Spreading wave of Ca2+ Fertilization envelope

10 sec after fertilization 25 sec 35 sec 1 min 500 µm Fig. 47-4a EXPERIMENT Figure 47.4 Is the distribution of Ca2+ in an egg correlated with formation of the fertilization envelope? 10 sec after fertilization 25 sec 35 sec 1 min 500 µm

1 sec before fertilization 10 sec after fertilization 20 sec 30 sec Fig. 47-4b RESULTS 1 sec before fertilization Figure 47.4 Is the distribution of Ca2+ in an egg correlated with formation of the fertilization envelope? 10 sec after fertilization 20 sec 30 sec 500 µm

CONCLUSION Point of sperm nucleus entry Spreading wave of Ca2+ Fig. 47-4c CONCLUSION Point of sperm nucleus entry Spreading wave of Ca2+ Fertilization envelope Figure 47.4 Is the distribution of Ca2+ in an egg correlated with formation of the fertilization envelope?

Activation of the Egg The sharp rise in Ca2+ in the egg’s cytosol increases the rates of cellular respiration and protein synthesis by the egg cell With these rapid changes in metabolism, the egg is said to be activated The sperm nucleus merges with the egg nucleus and cell division begins

Fertilization in Mammals Fertilization in mammals and other terrestrial animals is internal In mammalian fertilization, the cortical reaction modifies the zona pellucida, the extracellular matrix of the egg, as a slow block to polyspermy

Zona pellucida Follicle cell Sperm nucleus Cortical granules Fig. 47-5 Zona pellucida Follicle cell Figure 47.5 Fertilization in mammals Sperm nucleus Cortical granules Sperm basal body

In mammals the first cell division occurs 12–36 hours after sperm binding The diploid nucleus forms after this first division of the zygote

Cleavage Fertilization is followed by cleavage, a period of rapid cell division without growth Cleavage partitions the cytoplasm of one large cell into many smaller cells called blastomeres The blastula is a ball of cells with a fluid-filled cavity called a blastocoel

(a) Fertilized egg (b) Four-cell stage (c) Early blastula Fig. 47-6 (a) Fertilized egg Figure 47.6 Cleavage in an echinoderm embryo (b) Four-cell stage (c) Early blastula (d) Later blastula

(a) Fertilized egg Fig. 47-6a Figure 47.6 Cleavage in an echinoderm embryo (a) Fertilized egg

(b) Four-cell stage Fig. 47-6b Figure 47.6 Cleavage in an echinoderm embryo (b) Four-cell stage

(c) Early blastula Fig. 47-6c Figure 47.6 Cleavage in an echinoderm embryo (c) Early blastula

(d) Later blastula Fig. 47-6d Figure 47.6 Cleavage in an echinoderm embryo (d) Later blastula

The eggs and zygotes of many animals, except mammals, have a definite polarity The polarity is defined by distribution of yolk (stored nutrients) The vegetal pole has more yolk; the animal pole has less yolk

The three body axes are established by the egg’s polarity and by a cortical rotation following binding of the sperm Cortical rotation exposes a gray crescent opposite to the point of sperm entry

(a) The three axes of the fully developed embryo Fig. 47-7 Dorsal Right Anterior Posterior Left Ventral (a) The three axes of the fully developed embryo Animal pole Pigmented cortex First cleavage Animal hemisphere Point of sperm nucleus entry Figure 47.7 The body axes and their establishment in an amphibian Future dorsal side Vegetal hemisphere Gray crescent Vegetal pole (b) Establishing the axes

(a) The three axes of the fully developed embryo Fig. 47-7a Dorsal Right Anterior Posterior Left Ventral Figure 47.7 The body axes and their establishment in an amphibian (a) The three axes of the fully developed embryo

(b) Establishing the axes Fig. 47-7b-1 Animal pole Animal hemisphere Vegetal hemisphere Figure 47.7 The body axes and their establishment in an amphibian Vegetal pole (b) Establishing the axes

Point of sperm nucleus entry Fig. 47-7b-2 Pigmented cortex Point of sperm nucleus entry Future dorsal side Gray crescent Figure 47.7 The body axes and their establishment in an amphibian (b) Establishing the axes

(b) Establishing the axes Fig. 47-7b-3 First cleavage Figure 47.7 The body axes and their establishment in an amphibian (b) Establishing the axes

Point of sperm nucleus entry Animal hemisphere Fig. 47-7b-4 Animal pole First cleavage Pigmented cortex Point of sperm nucleus entry Animal hemisphere Future dorsal side Vegetal hemisphere Gray crescent Vegetal pole Figure 47.7 The body axes and their establishment in an amphibian (b) Establishing the axes

Cleavage planes usually follow a pattern that is relative to the zygote’s animal and vegetal poles

Zygote Fig. 47-8-1 Figure 47.8 Cleavage in a frog embryo For the Cell Biology Video Cleavage of a Fertilized Egg, go to Animation and Video Files.

Fig. 47-8-2 2-cell stage forming Figure 47.8 Cleavage in a frog embryo

Fig. 47-8-3 4-cell stage forming Figure 47.8 Cleavage in a frog embryo

Animal pole 8-cell stage Vegetal pole Fig. 47-8-4 Figure 47.8 Cleavage in a frog embryo

Blastula (cross section) Fig. 47-8-5 Blastocoel Blastula (cross section) Figure 47.8 Cleavage in a frog embryo

Blastula (cross section) Fig. 47-8-6 0.25 mm 0.25 mm Animal pole Blastocoel Figure 47.8 Cleavage in a frog embryo Vegetal pole Zygote 2-cell stage forming 4-cell stage forming 8-cell stage Blastula (cross section)

Cell division is slowed by yolk Holoblastic cleavage, complete division of the egg, occurs in species whose eggs have little or moderate amounts of yolk, such as sea urchins and frogs

Meroblastic cleavage, incomplete division of the egg, occurs in species with yolk-rich eggs, such as reptiles and birds

Gastrulation Gastrulation rearranges the cells of a blastula into a three-layered embryo, called a gastrula, which has a primitive gut

Video: Sea Urchin Embryonic Development The three layers produced by gastrulation are called embryonic germ layers The ectoderm forms the outer layer The endoderm lines the digestive tract The mesoderm partly fills the space between the endoderm and ectoderm Video: Sea Urchin Embryonic Development

Gastrulation in the sea urchin embryo The blastula consists of a single layer of cells surrounding the blastocoel Mesenchyme cells migrate from the vegetal pole into the blastocoel The vegetal plate forms from the remaining cells of the vegetal pole and buckles inward through invagination

Gastrulation in the sea urchin embryo The newly formed cavity is called the archenteron This opens through the blastopore, which will become the anus

Future ectoderm Future mesoderm Future endoderm Animal pole Blastocoel Fig. 47-9-1 Future ectoderm Future mesoderm Future endoderm Animal pole Blastocoel Mesenchyme cells Figure 47.9 Gastrulation in a sea urchin embryo Vegetal plate Vegetal pole

Future ectoderm Future mesoderm Future endoderm Fig. 47-9-2 Figure 47.9 Gastrulation in a sea urchin embryo

Filopodia pulling archenteron tip Fig. 47-9-3 Future ectoderm Future mesoderm Future endoderm Filopodia pulling archenteron tip Figure 47.9 Gastrulation in a sea urchin embryo Archenteron

Future ectoderm Future mesoderm Future endoderm Blastocoel Archenteron Fig. 47-9-4 Future ectoderm Future mesoderm Future endoderm Blastocoel Figure 47.9 Gastrulation in a sea urchin embryo Archenteron Blastopore

Mesenchyme (mesoderm forms future skeleton) Fig. 47-9-5 Future ectoderm Future mesoderm Future endoderm Ectoderm Mouth Figure 47.9 Gastrulation in a sea urchin embryo Mesenchyme (mesoderm forms future skeleton) Digestive tube (endoderm) Anus (from blastopore)

Figure 47.9 Gastrulation in a sea urchin embryo Key Future ectoderm Future mesoderm Future endoderm Archenteron Blastocoel Filopodia pulling archenteron tip Animal pole Blastocoel Archenteron Blastocoel Blastopore Mesenchyme cells Ectoderm Vegetal plate Vegetal pole Mouth Figure 47.9 Gastrulation in a sea urchin embryo Mesenchyme cells Mesenchyme (mesoderm forms future skeleton) Digestive tube (endoderm) Blastopore 50 µm Anus (from blastopore)

Gastrulation in the frog The frog blastula is many cell layers thick Cells of the dorsal lip originate in the gray crescent and invaginate to create the archenteron Cells continue to move from the embryo surface into the embryo by involution These cells become the endoderm and mesoderm

Gastrulation in the frog The blastopore encircles a yolk plug when gastrulation is completed The surface of the embryo is now ectoderm, the innermost layer is endoderm, and the middle layer is mesoderm

Fig. 47-10-1 SURFACE VIEW CROSS SECTION Animal pole Blastocoel Dorsal lip of blasto- pore Dorsal lip of blastopore Key Blastopore Future ectoderm Figure 47.10 Gastrulation in a frog embryo Early gastrula Future mesoderm Vegetal pole Future endoderm

Fig. 47-10-2 SURFACE VIEW CROSS SECTION Blastocoel shrinking Archenteron Key Future ectoderm Figure 47.10 Gastrulation in a frog embryo Future mesoderm Future endoderm

Fig. 47-10-3 SURFACE VIEW CROSS SECTION Ectoderm Mesoderm Blastocoel remnant Endoderm Archenteron Key Blastopore Future ectoderm Figure 47.10 Gastrulation in a frog embryo Late gastrula Future mesoderm Blastopore Yolk plug Future endoderm

Figure 47.10 Gastrulation in a frog embryo SURFACE VIEW CROSS SECTION Animal pole Blastocoel Dorsal lip of blasto- pore Dorsal lip of blastopore Blastopore Early gastrula Vegetal pole Blastocoel shrinking Archenteron Figure 47.10 Gastrulation in a frog embryo Ectoderm Mesoderm Blastocoel remnant Endoderm Archenteron Key Blastopore Future ectoderm Future mesoderm Late gastrula Blastopore Yolk plug Future endoderm

Gastrulation in the chick The embryo forms from a blastoderm and sits on top of a large yolk mass During gastrulation, the upper layer of the blastoderm (epiblast) moves toward the midline of the blastoderm and then into the embryo toward the yolk

The midline thickens and is called the primitive streak The movement of different epiblast cells gives rise to the endoderm, mesoderm, and ectoderm

Migrating cells (mesoderm) Hypoblast Fig. 47-11 Dorsal Fertilized egg Primitive streak Anterior Embryo Left Right Yolk Posterior Ventral Primitive streak Epiblast Future ectoderm Figure 47.11 Gastrulation in a chick embryo Blastocoel Endoderm Migrating cells (mesoderm) Hypoblast YOLK

Organogenesis During organogenesis, various regions of the germ layers develop into rudimentary organs The frog is used as a model for organogenesis

Early in vertebrate organogenesis, the notochord forms from mesoderm, and the neural plate forms from ectoderm

Figure 47.12 Early organogenesis in a frog embryo Neural folds Eye Somites Tail bud Neural fold Neural plate SEM 1 mm 1 mm Neural tube Neural crest cells Neural fold Neural plate Notochord Neural crest cells Coelom Somite Notochord Ectoderm Figure 47.12 Early organogenesis in a frog embryo Archenteron (digestive cavity) Mesoderm Outer layer of ectoderm Endoderm Neural crest cells (c) Somites Archenteron (a) Neural plate formation Neural tube (b) Neural tube formation

(a) Neural plate formation Fig. 47-12a Neural fold Neural plate 1 mm Neural folds Notochord Ectoderm Mesoderm Figure 47.12a Early organogenesis in a frog embryo Endoderm Archenteron (a) Neural plate formation

Video: Frog Embryo Development The neural plate soon curves inward, forming the neural tube The neural tube will become the central nervous system (brain and spinal cord) Video: Frog Embryo Development

(b) Neural tube formation Fig. 47-12b-1 Neural fold Neural plate Figure 47.12b Early organogenesis in a frog embryo (b) Neural tube formation

(b) Neural tube formation Fig. 47-12b-2 Figure 47.12b Early organogenesis in a frog embryo (b) Neural tube formation

(b) Neural tube formation Fig. 47-12b-3 Neural crest cells Figure 47.12b Early organogenesis in a frog embryo (b) Neural tube formation

Outer layer of ectoderm Fig. 47-12b-4 Outer layer of ectoderm Neural crest cells Neural tube Figure 47.12b Early organogenesis in a frog embryo (b) Neural tube formation

Neural crest cells develop along the neural tube of vertebrates and form various parts of the embryo (nerves, parts of teeth, skull bones, and so on) Mesoderm lateral to the notochord forms blocks called somites Lateral to the somites, the mesoderm splits to form the coelom

Archenteron (digestive cavity) Fig. 47-12c Neural tube Neural crest cells Eye Somites Tail bud Notochord Coelom Somite Archenteron (digestive cavity) Figure 47.12c Early organogenesis in a frog embryo SEM 1 mm (c) Somites

Organogenesis in the chick is quite similar to that in the frog

These layers form extraembryonic membranes Fig. 47-13 Eye Neural tube Notochord Forebrain Somite Coelom Heart Archenteron Endoderm Lateral fold Mesoderm Blood vessels Ectoderm Somites Yolk stalk Yolk sac These layers form extraembryonic membranes Figure 47.13 Organogenesis in a chick embryo Neural tube YOLK (a) Early organogenesis (b) Late organogenesis

These layers form extraembryonic membranes Fig. 47-13a Neural tube Notochord Somite Coelom Archenteron Endoderm Lateral fold Mesoderm Ectoderm Yolk stalk Figure 47.13 Organogenesis in a chick embryo Yolk sac These layers form extraembryonic membranes YOLK (a) Early organogenesis

(b) Late organogenesis Fig. 47-13b Eye Forebrain Heart Blood vessels Somites Figure 47.13 Organogenesis in a chick embryo Neural tube (b) Late organogenesis

The mechanisms of organogenesis in invertebrates are similar, but the body plan is very different

Reproductive system (except germ cells) Fig. 47-14 ECTODERM MESODERM ENDODERM Epidermis of skin and its derivatives (including sweat glands, hair follicles) Epithelial lining of mouth and anus Cornea and lens of eye Nervous system Sensory receptors in epidermis Adrenal medulla Tooth enamel Epithelium of pineal and pituitary glands Notochord Skeletal system Muscular system Muscular layer of stomach and intestine Excretory system Circulatory and lymphatic systems Reproductive system (except germ cells) Dermis of skin Lining of body cavity Adrenal cortex Epithelial lining of digestive tract Epithelial lining of respiratory system Lining of urethra, urinary bladder, and reproductive system Liver Pancreas Thymus Thyroid and parathyroid glands Figure 47.14 Adult derivatives of the three embryonic germ layers in vertebrates

Developmental Adaptations of Amniotes Embryos of birds, other reptiles, and mammals develop in a fluid-filled sac in a shell or the uterus Organisms with these adaptations are called amniotes

During amniote development, four extraembryonic membranes form around the embryo: The chorion functions in gas exchange The amnion encloses the amniotic fluid The yolk sac encloses the yolk The allantois disposes of waste products and contributes to gas exchange

Amniotic cavity with amniotic Fig. 47-15 Amnion Allantois Embryo Amniotic cavity with amniotic fluid Albumen Shell Figure 47.15 Extraembryonic membranes in birds and other reptiles Yolk (nutrients) Chorion Yolk sac

Mammalian Development The eggs of placental mammals Are small and store few nutrients Exhibit holoblastic cleavage Show no obvious polarity Gastrulation and organogenesis resemble the processes in birds and other reptiles Early cleavage is relatively slow in humans and other mammals

At completion of cleavage, the blastocyst forms A group of cells called the inner cell mass develops into the embryo and forms the extraembryonic membranes The trophoblast, the outer epithelium of the blastocyst, initiates implantation in the uterus, and the inner cell mass of the blastocyst forms a flat disk of cells As implantation is completed, gastrulation begins

Endometrial epithelium (uterine lining) Fig. 47-16-1 Endometrial epithelium (uterine lining) Uterus Inner cell mass Trophoblast Figure 47.16 Four stages in early embryonic development of a human Blastocoel

Epiblast Expanding region of trophoblast Maternal blood vessel Fig. 47-16-2 Expanding region of trophoblast Maternal blood vessel Epiblast Hypoblast Figure 47.16 Four stages in early embryonic development of a human Trophoblast

The epiblast cells invaginate through a primitive streak to form mesoderm and endoderm The placenta is formed from the trophoblast, mesodermal cells from the epiblast, and adjacent endometrial tissue The placenta allows for the exchange of materials between the mother and embryo By the end of gastrulation, the embryonic germ layers have formed

Epiblast Expanding region of trophoblast Amniotic cavity Hypoblast Fig. 47-16-3 Expanding region of trophoblast Amniotic cavity Epiblast Hypoblast Yolk sac (from hypoblast) Figure 47.16 Four stages in early embryonic development of a human Extraembryonic mesoderm cells (from epiblast) Chorion (from trophoblast)

Ectoderm Endoderm Amnion Chorion Mesoderm Yolk sac Fig. 47-16-4 Amnion Chorion Ectoderm Mesoderm Endoderm Yolk sac Figure 47.16 Four stages in early embryonic development of a human Extraembryonic mesoderm Atlantois

Figure 47.16 Four stages in early embryonic development of a human Endometrial epithelium (uterine lining) Expanding region of trophoblast Maternal blood vessel Uterus Inner cell mass Epiblast Trophoblast Hypoblast Blastocoel Trophoblast Expanding region of trophoblast Amnion Amniotic cavity Chorion Ectoderm Epiblast Mesoderm Figure 47.16 Four stages in early embryonic development of a human Hypoblast Endoderm Yolk sac (from hypoblast) Yolk sac Extraembryonic mesoderm cells (from epiblast) Extraembryonic mesoderm Chorion (from trophoblast) Allantois

The extraembryonic membranes in mammals are homologous to those of birds and other reptiles and develop in a similar way

Concept 47.2: Morphogenesis in animals involves specific changes in cell shape, position, and adhesion Morphogenesis is a major aspect of development in plants and animals Only in animals does it involve the movement of cells

The Cytoskeleton, Cell Motility, and Convergent Extension Changes in cell shape usually involve reorganization of the cytoskeleton Microtubules and microfilaments affect formation of the neural tube

Fig. 47-17-1 Ectoderm Figure 47.17 Change in cell shape during morphogenesis

Neural plate Microtubules Fig. 47-17-2 Figure 47.17 Change in cell shape during morphogenesis

Fig. 47-17-3 Actin filaments Figure 47.17 Change in cell shape during morphogenesis

Fig. 47-17-4 Figure 47.17 Change in cell shape during morphogenesis

Fig. 47-17-5 Neural tube Figure 47.17 Change in cell shape during morphogenesis

Ectoderm Neural plate Microtubules Actin filaments Neural tube Fig. 47-17-6 Ectoderm Neural plate Microtubules Actin filaments Figure 47.17 Change in cell shape during morphogenesis Neural tube

The cytoskeleton also drives cell migration, or cell crawling, the active movement of cells In gastrulation, tissue invagination is caused by changes in cell shape and migration Cell crawling is involved in convergent extension, a morphogenetic movement in which cells of a tissue become narrower and longer

Convergence Extension Fig. 47-18 Figure 47.18 Convergent extension of a sheet of cells For the Cell Biology Video Lamellipodia in Cell Migration, go to Animation and Video Files.

Role of Cell Adhesion Molecules and the Extracellular Matrix Cell adhesion molecules located on cell surfaces contribute to cell migration and stable tissue structure One class of cell-to-cell adhesion molecule is the cadherins, which are important in formation of the frog blastula

Embryo without EP cadherin Fig. 47-19 RESULTS 0.25 mm 0.25 mm Figure 47.19 Is cadherin required for development of the blastula? For the Cell Biology Video E-cadherin Expression, go to Animation and Video Files. Control embryo Embryo without EP cadherin

Fibers of the extracellular matrix may function as tracks, directing migrating cells along routes Several kinds of glycoproteins, including fibronectin, promote cell migration by providing molecular anchorage for moving cells

RESULTS Experiment 1 Control Matrix blocked Experiment 2 Control Fig. 47-20 RESULTS Experiment 1 Control Matrix blocked Experiment 2 Figure 47.20 Is an organized fibronectin matrix required for convergent extension? Control Matrix blocked

RESULTS Experiment 1 Control Matrix blocked Fig. 47-20-1 Figure 47.20 Is an organized fibronectin matrix required for convergent extension? Control Matrix blocked

RESULTS Experiment 2 Control Matrix blocked Fig. 47-20-2 Figure 47.20 Is an organized fibronectin matrix required for convergent extension? Control Matrix blocked

Concept 47.3: The developmental fate of cells depends on their history and on inductive signals Cells in a multicellular organism share the same genome Differences in cell types is the result of differentiation, the expression of different genes

Two general principles underlie differentiation: During early cleavage divisions, embryonic cells must become different from one another If the egg’s cytoplasm is heterogenous, dividing cells vary in the cytoplasmic determinants they contain

After cell asymmetries are set up, interactions among embryonic cells influence their fate, usually causing changes in gene expression This mechanism is called induction, and is mediated by diffusible chemicals or cell-cell interactions

Fate Mapping Fate maps are general territorial diagrams of embryonic development Classic studies using frogs indicated that cell lineage in germ layers is traceable to blastula cells

Blastomeres injected with dye Fig. 47-21 Epidermis Epidermis Central nervous system 64-cell embryos Notochord Blastomeres injected with dye Mesoderm Endoderm Blastula Neural tube stage (transverse section) Larvae Figure 47.21 Fate mapping for two chordates (a) Fate map of a frog embryo (b) Cell lineage analysis in a tunicate

Central nervous system Epidermis Fig. 47-21a Epidermis Central nervous system Epidermis Notochord Mesoderm Endoderm Figure 47.21a Fate mapping for two chordates Blastula Neural tube stage (transverse section) (a) Fate map of a frog embryo

Techniques in later studies marked an individual blastomere during cleavage and followed it through development

Blastomeres injected with dye Fig. 47-21b 64-cell embryos Blastomeres injected with dye Figure 47.21b Fate mapping for two chordates Larvae (b) Cell lineage analysis in a tunicate

Time after fertilization (hours) Fig. 47-22 Zygote First cell division Nervous system, outer skin, muscula- ture Muscula- ture, gonads Outer skin, nervous system Germ line (future gametes) Time after fertilization (hours) Musculature 10 Hatching Intestine Intestine Figure 47.22 Cell lineage in Caenorhabditis elegans Mouth Anus Eggs Vulva ANTERIOR POSTERIOR 1.2 mm

Establishing Cellular Asymmetries To understand how embryonic cells acquire their fates, think about how basic axes of the embryo are established

The Axes of the Basic Body Plan In nonamniotic vertebrates, basic instructions for establishing the body axes are set down early during oogenesis, or fertilization In amniotes, local environmental differences play the major role in establishing initial differences between cells and the body axes

Restriction of the Developmental Potential of Cells In many species that have cytoplasmic determinants, only the zygote is totipotent That is, only the zygote can develop into all the cell types in the adult

Unevenly distributed cytoplasmic determinants in the egg cell help establish the body axes These determinants set up differences in blastomeres resulting from cleavage

Experimental egg (side view) Fig. 47-23a EXPERIMENT Control egg (dorsal view) Experimental egg (side view) Gray crescent Gray crescent Thread Figure 47.23 How does distribution of the gray crescent affect the development potential of the first two daughter cells?

Experimental egg (side view) Fig. 47-23b EXPERIMENT Control egg (dorsal view) Experimental egg (side view) Gray crescent Gray crescent Thread Figure 47.23 How does distribution of the gray crescent affect the development potential of the first two daughter cells? RESULTS Normal Belly piece Normal

As embryonic development proceeds, potency of cells becomes more limited

Cell Fate Determination and Pattern Formation by Inductive Signals After embryonic cell division creates cells that differ from each other, the cells begin to influence each other’s fates by induction

The “Organizer” of Spemann and Mangold Based on their famous experiment, Hans Spemann and Hilde Mangold concluded that the blastopore’s dorsal lip is an organizer of the embryo The Spemann organizer initiates inductions that result in formation of the notochord, neural tube, and other organs

Dorsal lip of blastopore Primary embryo Fig. 47-24 EXPERIMENT RESULTS Dorsal lip of blastopore Primary embryo Secondary (induced) embryo Pigmented gastrula (donor embryo) Nonpigmented gastrula (recipient embryo) Primary structures: Neural tube Notochord Figure 47.24 Can the dorsal lip of the blastopore induce cells in another part of the amphibian embryo to change their developmental fate? Secondary structures: Notochord (pigmented cells) Neural tube (mostly nonpigmented cells)

Dorsal lip of blastopore Fig. 47-24a EXPERIMENT Dorsal lip of blastopore Pigmented gastrula (donor embryo) Figure 47.24 Can the dorsal lip of the blastopore induce cells in another part of the amphibian embryo to change their developmental fate? Nonpigmented gastrula (recipient embryo)

RESULTS Primary embryo Secondary (induced) embryo Fig. 47-24b RESULTS Primary embryo Secondary (induced) embryo Primary structures: Neural tube Figure 47.24 Can the dorsal lip of the blastopore induce cells in another part of the amphibian embryo to change their developmental fate? Notochord Secondary structures: Notochord (pigmented cells) Neural tube (mostly nonpigmented cells)

Formation of the Vertebrate Limb Inductive signals play a major role in pattern formation, development of spatial organization The molecular cues that control pattern formation are called positional information This information tells a cell where it is with respect to the body axes It determines how the cell and its descendents respond to future molecular signals

The wings and legs of chicks, like all vertebrate limbs, begin as bumps of tissue called limb buds

Figure 47.25 Vertebrate limb development Anterior Limb bud AER ZPA Limb buds Posterior 50 µm Apical ectodermal ridge (AER) (a) Organizer regions 2 Figure 47.25 Vertebrate limb development Digits 3 4 Anterior Ventral Proximal Distal Dorsal Posterior (b) Wing of chick embryo

Apical ectodermal ridge (AER) Fig. 47-25a Anterior Limb bud AER ZPA Limb buds Posterior 50 µm Apical ectodermal ridge (AER) Figure 47.25a Vertebrate limb development (a) Organizer regions

The embryonic cells in a limb bud respond to positional information indicating location along three axes Proximal-distal axis Anterior-posterior axis Dorsal-ventral axis

(b) Wing of chick embryo Fig. 47-25b 2 Digits 3 4 Anterior Ventral Proximal Distal Figure 47.25b Vertebrate limb development Dorsal Posterior (b) Wing of chick embryo

One limb-bud organizer region is the apical ectodermal ridge (AER) The AER is thickened ectoderm at the bud’s tip The second region is the zone of polarizing activity (ZPA) The ZPA is mesodermal tissue under the ectoderm where the posterior side of the bud is attached to the body

Tissue transplantation experiments support the hypothesis that the ZPA produces an inductive signal that conveys positional information indicating “posterior”

EXPERIMENT Anterior New ZPA Donor limb bud Host limb bud ZPA Posterior Fig. 47-26 EXPERIMENT Anterior New ZPA Donor limb bud Host limb bud ZPA Posterior RESULTS Figure 47.26 What role does the zone of polarizing activity (ZPA) play in limb pattern formation in vertebrates? 4 3 2 2 3 4

EXPERIMENT Anterior New ZPA Donor limb bud Host limb bud ZPA Posterior Fig. 47-26a EXPERIMENT Anterior New ZPA Donor limb bud Host limb bud Figure 47.26 What role does the zone of polarizing activity (ZPA) play in limb pattern formation in vertebrates? ZPA Posterior

Fig. 47-26b RESULTS 4 3 2 2 Figure 47.26 What role does the zone of polarizing activity (ZPA) play in limb pattern formation in vertebrates? 3 4

Signal molecules produced by inducing cells influence gene expression in cells receiving them Signal molecules lead to differentiation and the development of particular structures Hox genes also play roles during limb pattern formation

Fig. 47-27 Figure 47.27 Human polysyndactyly due to a homozygous mutation in a Hox gene

Cortical granule release (cortical reaction) Fig. 47-UN1 Sperm-egg fusion and depolarization of egg membrane (fast block to polyspermy) Cortical granule release (cortical reaction) Formation of fertilization envelope (slow block to polyspermy)

2-cell stage forming Animal pole 8-cell stage Vegetal pole Blastocoel Fig. 47-UN2 2-cell stage forming Animal pole 8-cell stage Vegetal pole Blastocoel Blastula

Fig. 47-UN3

Fig. 47-UN4 Neural tube Neural tube Notochord Notochord Coelom Coelom

Fig. 47-UN5 Species: Stage:

Fig. 47-UN6

You should now be able to: Describe the acrosomal reaction Describe the cortical reaction Distinguish among meroblastic cleavage and holoblastic cleavage Compare the formation of a blastula and gastrulation in a sea urchin, a frog, and a chick List and explain the functions of the extraembryonic membranes

Describe the process of convergent extension Describe the role of the extracellular matrix in embryonic development Describe two general principles that integrate our knowledge of the genetic and cellular mechanisms underlying differentiation Explain the significance of Spemann’s organizer in amphibian development

Explain pattern formation in a developing chick limb, including the roles of the apical ectodermal ridge and the zone of polarizing activity