LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane

LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane

LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 47 Animal Development Lectures by Erin Barley Kathleen Fitzpatrick 2011 Pearson Education, Inc. Overview: A Body-Building Plan A human embryo at about 7 weeks after conception shows development of distinctive

features 2011 Pearson Education, Inc. Figure 47.1 1 mm Development occurs at many points in the life cycle of an animal This includes metamorphosis and gamete production, as well as embryonic development 2011 Pearson Education, Inc. Figure 47.2

EMBRYONIC DEVELOPMENT Sperm FER TI Egg LIZ ATI O Adult frog

N Zygote EA L C Metamorphosis STR ULA TI NO

GA IS OR NES GE Larval stages GA GE A V Tail-bud embryo

Blastula ON Gastrula Although animals display different body plans, they share many basic mechanisms of development and use a common set of regulatory genes Biologists use model organisms to study development, chosen for the ease with which they can be studied in the laboratory 2011 Pearson Education, Inc. Concept 47.1: Fertilization and cleavage

initiate embryonic development Fertilization is the formation of a diploid zygote from a haploid egg and sperm 2011 Pearson Education, Inc. Fertilization Molecules and events at the egg surface play a crucial role in each step of fertilization Sperm penetrate the protective layer around the egg Receptors on the egg surface bind to molecules on the sperm surface Changes at the egg surface prevent polyspermy, the entry of multiple sperm nuclei into the egg

2011 Pearson Education, Inc. 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 2011 Pearson Education, Inc. Figure 47.3-5 Sperm plasma membrane

Sperm nucleus Basal body (centriole) Sperm head Acrosome Jelly coat Sperm-binding receptors Fertilization envelope Acrosomal

process Actin filament Cortical Fused granule plasma membranes Hydrolytic enzymes Perivitelline space Vitelline layer Egg plasma membrane

EGG CYTOPLASM Gamete contact and/or fusion depolarizes the egg cell membrane and sets up a fast block to polyspermy 2011 Pearson Education, Inc. The Cortical Reaction Fusion of egg and sperm also initiates the cortical reaction Seconds after the sperm binds to the egg, vesicles just beneath the egg plasma membrane release their contents and form a fertilization envelope The fertilization envelope acts as the slow block to polyspermy

2011 Pearson Education, Inc. The cortical reaction requires a high concentration of Ca2 ions in the egg The reaction is triggered by a change in Ca 2 concentration Ca2 spread across the egg correlates with the appearance of the fertilization envelope 2011 Pearson Education, Inc. Figure 47.4 EXPERIMENT

10 sec after fertilization 25 sec 35 sec 1 min 10 sec after fertilization 20 sec 30 sec

500 m RESULTS 1 sec before fertilization CONCLUSION Point of sperm nucleus entry Spreading wave of Ca2 Fertilization

envelope 500 m Egg Activation The rise in Ca2+ in the 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 proteins and mRNAs needed for activation are already present in the egg The sperm nucleus merges with the egg nucleus and cell division begins 2011 Pearson Education, Inc.

Fertilization in Mammals Fertilization in mammals and other terrestrial animals is internal Secretions in the mammalian female reproductive tract alter sperm motility and structure This is called capacitation, and must occur before sperm are able to fertilize an egg 2011 Pearson Education, Inc. Sperm travel through an outer layer of cells to reach the zona pellucida, the extracellular matrix of the egg When the sperm binds a receptor in the zona pellucida, it triggers a slow block to polyspermy

No fast block to polyspermy has been identified in mammals 2011 Pearson Education, Inc. Figure 47.5 Zona pellucida Follicle cell Sperm basal body Sperm nucleus

Cortical granules In mammals the first cell division occurs 1236 hours after sperm binding The diploid nucleus forms after this first division of the zygote 2011 Pearson Education, Inc. 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 2011 Pearson Education, Inc. Figure 47.6 50 m (a) Fertilized egg (b) Four-cell stage (c) Early blastula (d) Later blastula Cleavage Patterns In frogs and many other animals, the distribution of

yolk (stored nutrients) is a key factor influencing the pattern of cleavage The vegetal pole has more yolk; the animal pole has less yolk The difference in yolk distribution results in animal and vegetal hemispheres that differ in appearance 2011 Pearson Education, Inc. The first two cleavage furrows in the frog form four equally sized blastomeres The third cleavage is asymmetric, forming unequally sized blastomeres 2011 Pearson Education, Inc.

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 2011 Pearson Education, Inc. Figure 47.7 Zygote 2-cell stage

forming Gray crescent 0.25 mm 8-cell stage (viewed from the animal pole) 4-cell stage forming 8-cell stage Animal pole

0.25 mm Blastula (at least 128 cells) Vegetal pole Blastula (cross section) Blastocoel Regulation of Cleavage Animal embryos complete cleavage when the ratio of material in the nucleus relative to the cytoplasm is sufficiently large 2011 Pearson Education, Inc.

Concept 47.2: Morphogenesis in animals involves specific changes in cell shape, position, and survival After cleavage, the rate of cell division slows and the normal cell cycle is restored Morphogenesis, the process by which cells occupy their appropriate locations, involves Gastrulation, the movement of cells from the blastula surface to the interior of the embryo Organogenesis, the formation of organs 2011 Pearson Education, Inc. Gastrulation Gastrulation rearranges the cells of a blastula

into a three-layered embryo, called a gastrula 2011 Pearson Education, Inc. 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 Each germ layer contributes to specific structures in the adult animal 2011 Pearson Education, Inc.

Video: Sea Urchin Embryonic Development 2011 Pearson Education, Inc. Figure 47.8 ECTODERM (outer layer of embryo) Epidermis of skin and its derivatives (including sweat glands, hair follicles) Nervous and sensory systems Pituitary gland, adrenal medulla Jaws and teeth Germ cells MESODERM (middle layer of embryo) Skeletal and muscular systems Circulatory and lymphatic systems Excretory and reproductive systems (except germ cells)

Dermis of skin Adrenal cortex ENDODERM (inner layer of embryo) Epithelial lining of digestive tract and associated organs (liver, pancreas) Epithelial lining of respiratory, excretory, and reproductive tracts and ducts Thymus, thyroid, and parathyroid glands The newly formed cavity is called the archenteron This opens through the blastopore, which will become the anus 2011 Pearson Education, Inc.

Figure 47.9 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole Blastocoel Filopodia

Mesenchyme cells Blastopore Archenteron 50 m Blastocoel Ectoderm Key Future ectoderm Future mesoderm Future endoderm Mouth Mesenchyme

(mesoderm forms future skeleton) Archenteron Blastopore Digestive tube (endoderm) Anus (from blastopore) Gastrulation in Frogs Frog gastrulation begins when a group of cells on the dorsal side of the blastula begins to invaginate This forms a crease along the region where the gray crescent formed The part above the crease is called the dorsal lip of the blastopore

2011 Pearson Education, Inc. Cells continue to move from the embryo surface into the embryo by involution These cells become the endoderm and mesoderm Cells on the embryo surface will form the ectoderm 2011 Pearson Education, Inc. Figure 47.10 1

CROSS SECTION SURFACE VIEW Animal pole Blastocoel Dorsal lip of blastopore Early Vegetal pole gastrula Blastopore Blastocoel shrinking

2 3 Blastocoel remnant Dorsal lip of blastopore Archenteron Ectoderm Mesoderm Endoderm

Key Future ectoderm Future mesoderm Future endoderm Late gastrula Blastopore Blastopore Yolk plug Archenteron

Figure 47.10a 1 CROSS SECTION SURFACE VIEW Animal pole Blastocoel Key Future ectoderm Future mesoderm Future endoderm

Dorsal lip of blastopore Early Vegetal pole gastrula Blastopore Dorsal lip of blastopore Figure 47.10b

2 Key Future ectoderm Future mesoderm Future endoderm Blastocoel shrinking Archenteron Figure 47.10c

3 Key Future ectoderm Future mesoderm Future endoderm Late gastrula Blastopore Blastocoel remnant

Ectoderm Mesoderm Endoderm Blastopore Yolk plug Archenteron Gastrulation in Humans Human eggs have very little yolk A blastocyst is the human equivalent of the blastula The inner cell mass is a cluster of cells at one end of the blastocyst

The trophoblast is the outer epithelial layer of the blastocyst and does not contribute to the embryo, but instead initiates implantation 2011 Pearson Education, Inc. Following implantation, the trophoblast continues to expand and a set of extraembryonic membranes is formed These enclose specialized structures outside of the embryo Gastrulation involves the inward movement from the epiblast, through a primitive streak, similar to the chick embryo 2011 Pearson Education, Inc.

Figure 47.12 1 Blastocyst reaches uterus. Uterus Endometrial epithelium (uterine lining) Inner cell mass Trophoblast Blastocoel 2 Blastocyst implants (7 days after fertilization). Expanding region of

trophoblast Maternal blood vessel Epiblast Hypoblast Trophoblast 3 Extraembryonic membranes start to form (1011 days), and gastrulation begins (13 days). Expanding region of trophoblast

Amniotic cavity Epiblast Hypoblast Yolk sac (from hypoblast) Extraembryonic mesoderm cells (from epiblast) Chorion (from trophoblast) 4 Gastrulation has produced a three-layered embryo with four extraembryonic membranes. Amnion Chorion Ectoderm

Mesoderm Endoderm Yolk sac Extraembryonic mesoderm Allantois Developmental Adaptations of Amniotes The colonization of land by vertebrates was made possible only after the evolution of The shelled egg of birds and other reptiles as well as monotremes (egg-laying mammals) The uterus of marsupial and eutherian mammals 2011 Pearson Education, Inc. In both adaptations, embryos are surrounded

by fluid in a sac called the amnion This protects the embryo from desiccation and allows reproduction on dry land Mammals and reptiles including birds are called amniotes for this reason 2011 Pearson Education, Inc. The four extraembryonic membranes that 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 2011 Pearson Education, Inc. Organogenesis During organogenesis, various regions of the germ layers develop into rudimentary organs Early in vertebrate organogenesis, the notochord forms from mesoderm, and the neural plate forms from ectoderm 2011 Pearson Education, Inc.

Figure 47.13 Eye Neural folds Neural fold Tail bud Neural plate SEM 1 mm Neural fold

Somites Neural tube Neural plate Notochord Neural crest cells 1 mm Neural crest cells

Coelom Notochord Somite Ectoderm Mesoderm Endoderm Neural crest cells Outer layer of ectoderm Archenteron

(a) Neural plate formation Neural tube (b) Neural tube formation (c) Somites Archenteron (digestive cavity) The neural plate soon curves inward, forming the neural tube The neural tube will become the central nervous system (brain and spinal cord)

2011 Pearson Education, Inc. Video: Frog Embryo Development 2011 Pearson Education, Inc. Figure 47.13b-3 Neural fold Neural plate Neural crest cells

Neural crest cells (b) Neural tube formation Neural tube Outer layer of ectoderm 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 (body cavity) 2011 Pearson Education, Inc. Organogenesis in the chick is quite similar to that in the frog 2011 Pearson Education, Inc. Figure 47.14 Neural tube Notochord

Eye Forebrain Somite Coelom Endoderm Mesoderm Ectoderm Archenteron Lateral fold Heart Blood vessels

Somites Yolk stalk These layers form extraembryonic membranes. (a) Early organogenesis Yolk sac Neural tube YOLK (b) Late organogenesis The mechanisms of organogenesis in

invertebrates are similar, but the body plan is very different For example, the neural tube develops along the ventral side of the embryo in invertebrats, rather than dorsally as occurs in vertebrates 2011 Pearson Education, Inc. Mechanisms of Morphogenesis Morphogenesis in animals but not plants involves movement of cells 2011 Pearson Education, Inc. The Cytoskeleton in Morphogenesis Reorganization of the cytoskeleton is a major force

in changing cell shape during development For example, in neurulation, microtubules oriented from dorsal to ventral in a sheet of ectodermal cells help lengthen the cells along that axis 2011 Pearson Education, Inc. Figure 47.15-5 Ectoderm Neural plate Microtubules Actin

filaments Neural tube The cytoskeleton promotes elongation of the archenteron in the sea urchin embryo This is convergent extension, the rearrangement of cells of a tissue that cause it to become narrower (converge) and longer (extend) Convergent extension occurs in other developmental processes The cytoskeleton also directs cell migration 2011 Pearson Education, Inc. Figure 47.16

Co ce n e g r e nv Exten sio n

Programmed Cell Death Programmed cell death is also called apoptosis At various times during development, individual cells, sets of cells, or whole tissues stop developing and are engulfed by neighboring cells For example, many more neurons are produced in developing embryos than will be needed Extra neurons are removed by apoptosis 2011 Pearson Education, Inc. Concept 47.3: Cytoplasmic determinants and inductive signals contribute to cell fate specification Determination is the term used to describe the process by which a cell or group of cells becomes

committed to a particular fate Differentiation refers to the resulting specialization in structure and function 2011 Pearson Education, Inc. Cells in a multicellular organism share the same genome Differences in cell types is the result of the expression of different sets of genes 2011 Pearson Education, Inc. Fate Mapping Fate maps are diagrams showing organs and other structures that arise from each region of an

embryo Classic studies using frogs indicated that cell lineage in germ layers is traceable to blastula cells 2011 Pearson Education, Inc. Figure 47.17 Epidermis Central nervous system Notochord Epidermis

Mesoderm Endoderm Blastula Neural tube stage (transverse section) (a) Fate map of a frog embryo 64-cell embryos Blastomeres injected with dye Larvae

(b) Cell lineage analysis in a tunicate Figure 47.17a Epidermis Central nervous system Notochord Epidermis Mesoderm Endoderm Blastula

(a) Fate map of a frog embryo Neural tube stage (transverse section) Figure 47.17b 64-cell embryos Blastomeres injected with dye Larvae (b) Cell lineage analysis in a tunicate Later studies of C. elegans used the ablation

(destruction) of single cells to determine the structures that normally arise from each cell The researchers were able to determine the lineage of each of the 959 somatic cells in the worm 2011 Pearson Education, Inc. Time after fertilization (hours) Figure 47.18 Zygote 0

First cell division Nervous system, outer skin, musculature 10 Outer skin, nervous system Musculature, gonads Germ line (future gametes)

Musculature Hatching Intestine Intestine Anus Mouth Eggs ANTERIOR Vulva 1.2 mm

POSTERIOR Germ cells are the specialized cells that give rise to sperm or eggs Complexes of RNA and protein are involved in the specification of germ cell fate In C. elegans, such complexes are called P granules, persist throughout development, and can be detected in germ cells of the adult worm 2011 Pearson Education, Inc. Figure 47.19 100 m

P granules are distributed throughout the newly fertilized egg and move to the posterior end before the first cleavage division With each subsequent cleavage, the P granules are partitioned into the posterior-most cells P granules act as cytoplasmic determinants, fixing germ cell fate at the earliest stage of development 2011 Pearson Education, Inc. Figure 47.20 20 m 1 Newly fertilized egg

2 Zygote prior to first division 3 Two-cell embryo 4 Four-cell embryo Axis Formation A body plan with bilateral symmetry is found across a range of animals This body plan exhibits asymmetry across the dorsal-ventral and anterior-posterior axes The right-left axis is largely symmetrical 2011 Pearson Education, Inc.

The anterior-posterior axis of the frog embryo is determined during oogenesis The animal-vegetal asymmetry indicates where the anterior-posterior axis forms The dorsal-ventral axis is not determined until fertilization 2011 Pearson Education, Inc. Upon fusion of the egg and sperm, the egg surface rotates with respect to the inner cytoplasm This cortical rotation brings molecules from one area of the inner cytoplasm of the animal hemisphere to interact with molecules in the vegetal cortex This leads to expression of dorsal- and ventralspecific gene expression

2011 Pearson Education, Inc. Figure 47.21 Dorsal Right Anterior Posterior Left Ventral (a) The three axes of the fully developed embryo Animal pole

Animal hemisphere Vegetal hemisphere Vegetal pole (b) Establishing the axes Point of sperm nucleus entry Gray crescent

Pigmented cortex Future dorsal side First cleavage In chicks, gravity is involved in establishing the anterior-posterior axis Later, pH differences between the two sides of the blastoderm establish the dorsal-ventral axis In mammals, experiments suggest that orientation of the egg and sperm nuclei before fusion may

help establish embryonic axes 2011 Pearson Education, Inc. Restricting Developmental Potential Hans Spemann performed experiments to determine a cells developmental potential (range of structures to which it can give rise) Embryonic fates are affected by distribution of determinants and the pattern of cleavage The first two blastomeres of the frog embryo are totipotent (can develop into all the possible cell types) 2011 Pearson Education, Inc.

Figure 47.22-2 EXPERIMENT Control egg (dorsal view) Experimental egg (side view) 1a Control 1b Experimental group group

Gray crescent Gray crescent Thread 2 RESULTS Normal Belly piece

Normal In mammals, embryonic cells remain totipotent until the 8-cell stage, much longer than other organisms Progressive restriction of developmental potential is a general feature of development in all animals In general tissue-specific fates of cells are fixed by the late gastrula stage 2011 Pearson Education, Inc. Cell Fate Determination and Pattern Formation by Inductive Signals As embryonic cells acquire distinct fates, they

influence each others fates by induction 2011 Pearson Education, Inc. The Organizer of Spemann and Mangold Spemann and Mangold transplanted tissues between early gastrulas and found that the transplanted dorsal lip triggered a second gastrulation in the host The dorsal lip functions as an organizer of the embryo body plan, inducing changes in surrounding tissues to form notochord, neural tube, and so on 2011 Pearson Education, Inc.

Figure 47.23 EXPERIMENT Dorsal lip of blastopore Pigmented gastrula (donor embryo) RESULTS Primary embryo Secondary (induced) embryo Nonpigmented

gastrula (recipient embryo) Primary structures: Neural tube 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 2011 Pearson Education, Inc. The wings and legs of chicks, like all vertebrate limbs, begin as bumps of tissue called limb buds 2011 Pearson Education, Inc. Figure 47.24 Anterior Limb bud

AER ZPA Posterior Limb buds 2 50 m Digits Apical ectodermal ridge (AER)

Anterior 3 4 Ventral Proximal Distal Dorsal Posterior (a) Organizer regions

(b) Wing of chick embryo 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 2011 Pearson Education, Inc. One limb-bud regulating region is the apical ectodermal ridge (AER) The AER is thickened ectoderm at the buds 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 2011 Pearson Education, Inc. Tissue transplantation experiments support the hypothesis that the ZPA produces an inductive signal that conveys positional information indicating posterior 2011 Pearson Education, Inc. Figure 47.25 EXPERIMENT

Anterior New ZPA Donor limb bud Host limb bud ZPA Posterior RESULTS

4 3 2 2 4 3 Sonic hedgehog is an inductive signal for limb development Hox genes also play roles during limb pattern formation

2011 Pearson Education, Inc. Cilia and Cell Fate Ciliary function is essential for proper specification of cell fate in the human embryo Motile cilia play roles in left-right specification Monocilia (nonmotile cilia) play roles in normal kidney development 2011 Pearson Education, Inc.

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