Transcript Chapter 54

CHAPTER 54
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Fertilization
• In all sexually-reproducing animals, the
first step is fertilization – union of male and
female gametes
• Fertilization itself consists of three events
– Sperm penetration and membrane fusion
– Egg activation
– Fusion of nuclei
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Fertilization
• Sperm penetration and membrane fusion
– Protective layers of egg include the jelly layer
and vitelline envelope in sea urchins, and the
zona pellucida in mammals
– Sperm’s acrosome contains digestive
enzymes that enable the sperm to tunnel its
way through to the egg’s cell membrane
– Membrane fusion permits sperm nucleus to
enter directly into egg’s cytoplasm
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Sperm
Jelly layer
Plasma
membrane
Vitelline
envelope
Cytoplasm
Cortical granules
a.
Nucleus
of egg
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Granulosa cell
Sperm
Zona pellucida
Oocyte
Plasma
membrane
Granulosa
cell
First
polar
body
Cortical granules
c.
1.2 µm
Cytoplasm
b.
d.
c-d: © David M. Phillips/Visuals Unlimited
3.3 µm
Fertilization
1. Sperm penetrates
2. Some of the zona
4. The sperm nucleus
3. Sperm and egg
between granulosa
pellucida is degraded
dissociates and
plasma membranes
cells.
by acrosomal enzymes.
enters cytoplasm.
fuse.
Plasma
membrane
Granulosa
cells
Zona
pellucida
Cortical
granules
6. Additional sperm
can no longer penetrate
the zona pellucida.
5. Cortical granules
release enzymes that
harden zona pellucida
and strip it of sperm
receptors. Hyalin
attracts water by osmosis.
7. Sperm and egg
pronuclei are
enclosed in a
nuclear envelope.
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Fertilization
• Membrane fusion
– Egg activation
• Dramatic increase in the levels of free intracellular
Ca2+ ions in the egg shortly after the sperm makes
contact with the egg’s plasma membrane
• Act as second messengers to initiate changes
– Block to polyspermy
• Rapid transient change in membrane potential
• Cortical granules remove sperm receptors
• Vitelline envelope lifts off – fertilization envelope
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Fertilization
• Sperm penetration has three other effects
– Triggers the egg to complete meiosis
– Triggers a cytoplasmic rearrangement
– Causes a sharp increase in protein synthesis and
metabolic activity in general
Primary Oocyte
First Metaphase of Meiosis Second Metaphase of Meiosis
Diploid nucleus
Meiosis Complete
Polar
bodies
Polar
body
Female
pronucleus
(haploid)
• Roundworms (Ascaris)
• Polychaete worms (Myzostoma)
• Clam worms (Nereis)
• Clams (Spisula)
• Nemertean worms (Cerebratulus)
• Polychaete worms (Chaetopterus)
• Mollusks (Dentalium)
• Many insects
• Sea stars
• Lancelets (Branchiostoma)
• Amphibians
• Mammals
• Fish
• Cnidarians
• Sea urchins
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Fertilization
• Fusion of nuclei
– 3rd and final stage of fertilization
– Haploid sperm and haploid egg nuclei fuse to
form diploid nucleus of the zygote
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Cleavage
• Rapid division of the zygote into a larger
and larger number of smaller and smaller
cells (blastomeres)
• Not accompanied by an increase in the
overall size of the embryo
• Animal pole
– Forms external tissues
• Vegetal pole
– Forms internal tissues
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Cleavage
• Blastula
– Hollow ball of cells
– Blastocoel – fluid-filled cavity
• Cleavage patterns are quite diverse
– Relative amount of nutritive yolk in the egg is
the characteristic that most affects the
cleavage pattern of an animal embryo
– Vertebrates exhibit a variety of reproductive
strategies involving different patterns of yolk
utilization
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Cleavage Patterns
• Eggs with little or no yolk
– Holoblastic cleavage
– Invertebrates, amphibians, mammals
• Eggs with large amounts of yolk
– Meroblastic cleavage
– Embryo forms thin cap on yolk
Sea Urchin
Frog
Chicken
Animal pole
Nucleus
Cytoplasm
Cytoplasm
Shell
Nucleus
Air
bubble
Nucleus
Plasma
membrane
Albumen
Yolk
Yolk
Vegetal pole
a.
b.
Yolk
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c.
Holoblastic cleavage
Meroblastic cleavage
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Cleavage Patterns
• Mammalian eggs contain very little yolk
– Undergo holoblastic cleavage
– Form a blastocyst composed of
• Trophoblast
– Outer layer of cells
– Contributes to the placenta
• Blastocoel
– Central fluid-filled cavity
• Inner cell mass
– Located at one pole
– Forms the developing embryo
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Fate of Blastomeres
• In many animals removal of committed cells
results in embryos deficient in tissues that would
have developed from those tissues
• In mammals, early blastomeres do not appear to
be committed to a particular fate
– Cell removed for preimplantation genetic diagnosis
– Split embryos form identical twins
• In mammals, body form determined primarily by
cell-to-cell interactions
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Gastrulation
• Process involving a complex series of cell shape
changes and cell movements that occurs in the
blastula
• Establishes the basic body plan and creates the
three primary germ layers
– Ectoderm – Exterior
• Epidermis of skin, nervous system, sense organs
– Mesoderm – Middle
• Skeleton, muscles, blood vessels, heart, blood, gonads,
kidneys, dermis of skin
– Endoderm – Inner
• Lining of digestive and respiratory tracts, liver, pancreas,
thymus, thyroid
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Gastrulation
• Cells move during gastrulation using a
variety of cell shape changes
– Cells that are tightly attached to each other
via junctions will move as cell sheets
– Invagination – Cell sheet dents inward
– Involution – Cell sheet rolls inward
– Delamination – Cell sheet splits in two
– Ingression – Cells break away from cell sheet
and migrate as individual cells
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Gastrulation Patterns
• Vary according to the amount of yolk
• Gastrulation in sea urchins
– Develop from relatively yolk-poor eggs
– Form hollow symmetrical blastulas
– Deuterostome – anus develops first and
mouth second
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Gastrulation Patterns
• Gastrulation in mammals
– Proceeds similarly to that in birds
– Embryo develops from inner cell mass
– Embryo gastrulates as if it was sitting in a ball
of yolk
• Embryo obtains nutrition from placenta
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Extraembryonic Membranes
• Adaptation to life on dry land
– Reptiles, birds, and mammals
– Amniotic species developed
• Extraembryonic membranes
– Amnion, chorion, yolk sac, and allantois
• Nourish and protect the developing
embryo
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Extraembryonic Membranes
• Amnion
– Encloses amniotic fluid
• Chorion
– Located near eggshell in birds
– Contributes to the placenta in mammals
• Yolk sac
– Food source in bird embryos
– Found in mammals, but it is not nutritive
• Allantois
– Unites with chorion in birds, forming a structure used for gas
exchange
– In mammals, it contributes blood vessels to the developing
umbilical cord
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Extraembryonic Membranes
Chick Embryo
Mammal Embryo
Chorion
Amnion
Chorion
Yolk sac
Amnion
Umbilical
blood
vessels
Yolk sac
Villus of chorion
frondosum
Allantois
Maternal blood
a.
b.
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Organogenesis
• Formation of organs in their proper
locations
• Occurs by interaction of cells within and
between the three germ layers
• Thus, it follows rapidly on the heels of
gastrulation
– In many animals it begins before gastrulation
is complete
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Organogenesis
• To a large degree, a cell’s location in the
developing embryo determines its fate
• At some stage, every cell’s ultimate fate
becomes fixed – cell determination
• A cell’s fate can be established by
– Inheritance of cytoplasmic determinants
– Interactions with neighboring cells
• Induction
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Organogenesis in Vertebrates
• Begins with the formation of two structures
unique to chordates
– Notochord
– Dorsal nerve cord – neurulation
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Development of Neural Tube
• Notochord
– Forms from mesoderm
– Region of dorsal ectodermal cells situated
above notochord thickens to form the neural
plate
– Cells of the neural plate fold together to form
a long hollow cylinder, the neural tube
– Will become brain and spinal cord
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Neural plate
Amniotic
cavity
Ectoderm
Mesoderm
Notochord
Endoderm
Yolk sac
a.
Neural groove
Neural fold
Ectoderm
Notochord
Mesoderm
Endoderm
b.
Neural tube
Ectoderm
Neural crest
Mesoderm
Endoderm
Somite
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c.
Human Development
• Human development from fertilization to
birth takes an average of 266 days, or
about 9 months
– Commonly divided into three periods called
trimesters
28
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or display.
First Trimester
• First month
a.
© Lennart Nilsson/Albert Bonniers Förlag AB, A Child Is Born, Dell Publishing
Company
– Zygote undergoes its first cleavage about 30
hr after fertilization
– By the time the embryo reaches the uterus,
6–7 days after fertilization, it has differentiated
into a blastocyst
– Trophoblast cells digest their way into the
endometrium in the process known as
implantation
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First Trimester
• First month
– During the second week, the developing chorion and
mother’s endometrium engage to form the placenta
• Mom and baby’s blood come into close proximity, but do not
mix – gases are exchanged
• One hormone released by the placenta is human chorionic
gonadotropin (hCG)
–
–
–
–
Gastrulation occurs in the second week
Neurulation occurs in the third week
Organogenesis begins in the fourth week
Embryo is 5 mm in length
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Chorion
Amnion
Yolk sac
Umbilical
cord
Chorionic
frondosum
(fetal)
Decidua
basalis
(maternal)
Placenta
Umbilical artery
Umbilical vein
Uterine wall
a.
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reproduction or display.
First Trimester
• Second month
– Organogenesis continues
– Miniature limbs assume adult shape
– All major organs in the body established
– Embryo grows to about 25 mm in length
– Weighs about 1 g, and looks distinctly human
– 9th week marks the transition from embryo to
fetus
b.
© Lennart Nilsson/Albert Bonniers Förlag AB, A Child Is Born, Dell Publishing
Company
32
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required for reproduction or display.
First Trimester
• Third month
– Nervous system develops
c.
– Limbs start to move
– Secretion of hCG by the placenta declines,
and so corpus luteum degenerates
– Placenta takes over hormone secretion
© Lennart Nilsson/Albert Bonniers Förlag AB, A Child Is
Born, Dell Publishing Company
33
Increasing Hormone Concentration
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hCG
Estrogen
Progesterone
0
1
2
3
4
5
6
Months of Pregnancy
7
8
9
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reproduction or display.
Second Trimester
d.
© Lennart Nilsson/Albert Bonniers Förlag AB, A Child Is Born, Dell
Publishing Company
• The basic body plan develops further
• Bones actively enlarge in fourth month
• Rapid fetal heartbeat can be heard by a
stethoscope
• By the end of the sixth month, fetus is over
300 mm long, and weighs 600 g
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Third Trimester
• A period of growth and organ maturation
• Weight of the fetus doubles several times
• Most of the major nerve tracts in the brain
are formed
• Brain continues to develop and produce
neurons for months after birth
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Birth
• Estrogen stimulates mother’s uterus to
release prostaglandins, and produce more
oxytocin receptors
– Prostaglandins begin uterine contractions
– Sensory feedback from uterus stimulates
oxytocin release from posterior pituitary
• Oxytocin and prostaglandins further
stimulate uterine contractions
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Birth
• Strong contractions, aided by the mother’s
voluntary pushing, expel the fetus
• Now called a newborn baby, or neonate
• After birth, continuing uterine contractions
expel the placenta and associated
membranes
– Collectively called the afterbirth
38
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Intestine
Placenta
Umbilical
cord
Wall of
uterus
Cervix
Vagina
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Nursing
• Milk production (lactation) occurs in alveoli
of mammary glands when stimulated by
the anterior pituitary hormone prolactin
• During pregnancy, the mammary glands
are prepared for, but prevented from,
lactating
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Nursing
• After birth
– Prolactin – stimulates the mammary alveoli to
produce milk
– Suckling triggers posterior pituitary to release
oxytocin
• Stimulates contraction of smooth muscles
surrounding alveolar ducts
• Milk is ejected (milk let-down reflex)
• The first milk produced after birth, colostrum, is
rich in nutrients and maternal antibodies
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Postnatal Development
• Growth of the infant continues rapidly after
birth
• Babies typically double their birth weight
within 2 months
• Different components grow at different
rates
– Allometric growth
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