recapitulation theory / Haeckel's
biogenetic law : ontogeny recapitulates phylogeny;
that is, an organism in the course of its development goes through the
same successive stages as did the species in developing from the lower
to the higher forms of animal life
propagule : cell unit capable of developing
into a complete organism
morphogen : a diffusible substance in embryonic tissue postulated
to form a concentration gradient that influences morphogenesis.
morphogen gradients : hypotheses
on the development
free diffusion
argosomes (vesicle-mediated gradients) : Wnt and Hedgehog family
proteins are secreted signalling molecules (morphogens) that act at both
long and short range to control growth and patterning during development.
Both proteins are covalently modified by lipid, and the mechanism by which
such hydrophobic molecules might spread over long distances is unknown.
Wingless, Hedgehog and glycophosphatidylinositol-linked proteins copurify
with lipoprotein particles, and co-localize with them in the developing
wing epithelium of Drosophila. In larvae with reduced lipoprotein
levels, Hedgehog accumulates near its site of production, and fails to
signal over its normal range. Similarly, the range of Wingless signalling
is narrowed. A novel function for lipoprotein particles, in which they
act as vehicles for the movement of lipid-linked morphogens and glycophosphatidylinositol-linked
proteinsref
a cell's perception of morphogen concentration is thought to be determined
by the number of active receptors, with inactive receptors making little
if any contribution. Patched (Ptc),
the receptor for the morphogen Hedgehog
(Hh),
is active in the absence of ligand and blocks the expression of target
genes by inhibiting Smoothened (Smo), an essential transducer of the Hh
signal. Hh binding to Ptc abrogates the ability of Ptc to inhibit Smo,
thereby unleashing Smo activity and inducing target gene expression. A
cell's measure of ambient Hh concentration is not determined solely by
the number of active (unliganded) Ptc molecules. Instead, we find that
Hh-bound Ptc can titrate the inhibitory action of unbound Ptc. This effect
is sufficient to allow normal reading of the Hh gradient in the presence
of a form of Ptc that cannot bind the ligand but retains its ability to
inhibit Smo. These results support a model in which the ratio of bound
to unbound Ptc molecules determines the cellular response to Hhref.
ontogeny : the development of the individual
organism
morphogenetic movement : a flowing of cell groups concerned with
the formation of germ layers or of organ primordia
convergent extension (CE) of
the axial mesoderm of vertebrates is an intensively studied model for morphogenetic
processes that rely on cell rearrangement. It involves the intercalation
of polarized cells perpendicular to the antero-posterior (AP) axis, which
narrows and lengthens the tissue. The chordamesoderm of Xenopus
possesses an intrinsic AP polarity that is necessary for CE, functions
in parallel to Wnt/planar cell polarity signalling, and determines the
direction of tissue elongation. The mechanism that establishes AP polarity
involves graded activin-like signalling and directly links mesoderm AP
patterning to CEref
morphodifferentiation : the arrangement of formative cells in the
development of tissues or organs, which leads to production of the ultimate
shape of the structure.
cytomorphosis : the series of changes through which cells go in
the process of formation, development, senescence, etc.
epithelial stem
cell
differentiation : the first step is the alteration of the adhesive
properties of an individual cell. This is achieved by systematically changing
the complement of surface adhesion molecules used to attach to its neighbors.
Epithelial
buds are small bulges in undifferentiated embryonic epithelial layers
that develop into a variety of complex structures, including teeth, limbs,
and hair follicles. The formation of the epithelial bud is caused by a
decrease in E-cadherin
and an increase in P-cadherin
levels : only through combined noggin
and Wnt3a
signaling can the Lef1
transcription complex repress E-cadherin
expression, an intriguing outcome since Lef1 complexes typically activate
transcription in response to Wnt signaling.
emancipation : the establishment of local autonomy within restricted
fields of a developing embryo.
presumptive region : an area of the blastula which has been proved
under normal conditions to develop into a specific organ or type of tissue.
morphometry : the measurement of the forms or structures of organisms
morphon : an individual organism or structural unit.
morphophysics : the study of the physical and chemical causes of
development.
morphosis : the process of formation of a part or organ.
morphogenesia / morphogenesis :
the evolution and development of form, as the development of the shape
of a particular organ or part of the body, or the development undergone
by individuals who attain the type to which the majority of the individuals
of the species approximate. The developmental cascade of pattern formation,
body plan establishment, and the architecture of mirror-image bilateral
symmetry of many structures and asymmetry of some, culminating in the adult
form.
inductor : a tissue elaborating a chemical
substance which acts to determine the growth and differentiation of embryonic
parts
activator : a substance that stimulates
the development of a particular structure in the embryo
organizer : a part of an embryo which so
influences some other part as to bring about and direct its histological
and morphological differentiation. Parts developing as a result of induction,
and inducing in their turn are classified as organizers of the second grade,
third grade, and so on
primary organizer : the dorsal lip region of the blastopore.
dorsal lip region : the mesodermal tissue around the dorsal lip
of the blastopore of an amphibian; an analogous
region in the human embryo is the organizer which by induction initiates
and controls early development.
secondary organizer : one of second grade, such as the optic cup,
which exerts influence on the developing lens.
tertiary organizer : one of third grade, such as the tympanic ring,
which exerts influence on the tympanic membrane
defect experiment : observation of an embryo, after destruction
of a region or part, to ascertain the effect on development.
capacitation : the process by which spermatozoa
become capable of fertilizing an oocyte
(ovum)
after it reaches the ampullary portion of the uterine tube; a series of
biochemical transformations the spermatozoa undergo in the female reproductive
tract. Superoxide anion production output by spermatozoa, hydroxyl radical
and hydrogen peroxide initiate sperm capacitation. Cholesterol efflux
=> elevation of intracellular Ca2+ and HCO3-
=> activation of adenylyl cyclase (AC) => cAMP => activates PKA to indirectly
phosphorylate certain proteins on tyrosine. Uncertainties exist as to the
number of forms of AC that are present in the spermatozoon, whether they
are cytosolic or bound to subcellular structures, and which physiological
effectors they respond to (e.g. bicarbonate, Ca2+, or receptor-coupled
G-proteins). While net intracellular levels of cAMP in spermatozoa depend
upon the relative activities of AC and PDE, there are wide between-sample
variations within species, both in basal levels and in levels attained
after activation of the AC (e.g. after sperm treatment with HCO3-).
Moreover, minor changes in bulk cAMP levels can result in large changes
in cAMP-dependent functions. Finally, while cAMP levels respond very rapidly
to sperm treatment by effectors such as HCO3- and
Ca2+ (key components of capacitating media), there are big discrepancies
between the rates of functional response. For example, enhancement of motility
and collapse of phospholipid asymmetry take place within a few minutes,
whereas more than 1 h of exposure to capacitating conditions is needed
for cAMP-dependent protein tyrosine phosphorylation to become detectable
or for the sperm population to attain a capacitated state. During capacitation,
there is also an increase in protein tyrosine phosphorylation dependent
actin polymerization and in the membrane-bound PLC. The main tyrosine phosphorylated
proteins during the course of capacitation and fertilization are localized
to the flagellum, although tyrosine phosphorylation of less abundant proteins
may also be regulated in the sperm head. Spermatozoa bound to the zona
pellucida and fusing with the oocyte plasma membrane are characterized
by a tyrosine phosphorylated flagellum.
sperm
hyperactivation => hyperactivated motility : the movement of hyperactivated
sperm appears different under different physical conditions and in different
species, but basically it involves an increase in flagellar bend amplitude
and, usually, beat asymmetry. Hyperactivation may be critical to the success
of fertilization, because it enhances the ability of sperm to detach from
the wall of the oviduct, to move around in the labyrinthine lumen of the
oviduct, to penetrate mucous substances and, finally, to penetrate the
zona pellucida of the oocyte. Presumably, a signal or signals exist in
the oviduct to initiate hyperactivation at the appropriate time; however,
none has yet been identified with certainty. There is evidence that the
source of the signal is follicular fluid, yet spermatozoa are known to
hyperactivate before ovulation would release the fluid into the oviduct.
It requires sperm-specific voltage-gated calcium channels CatSper1
and CatSper2
in the flagellum => calcium ions interact with the axoneme of the flagellum
=> protein phosphorylation in the flagellum => switch on hyperactivation.
The process may also involve increases in intracellular cAMP, which at
least is required to support motility in general. The edding of antioxidant
enzymes inhibits the spontaneous and induced sperm hyperactivation : anyway
high doses of reactive oxygen species block the sperm motility through
the inhibition of ATP synthesis by the mitochondrial enzymes and cell membrane
compounds injury. Although hyperactivation often occurs during the process
of capacitation, divergent pathways regulate the 2 events. Sperm hyperactivation
might be a moiety of capacitation or the result of capacitation.
conceptus : the sum of derivatives of a
fertilized oocyte (ovum) at any stage of development from fertilization
until birth, including extraembryonic membranes as well as the embryo or
fetus
gestational age (GA) : age of conceptus
or pregnancy.
In human clinical practice, pregnancy is timed from onset of the last normal
menstruation and lasts 40 weeks (280 days). Elsewhere the onset may be
timed from estrus, coitus, artificial insemination, vaginal plug formation,
fertilization, or implantation.
menstrual age : conceptus age defined
by the time elapsed since the onset of the mother's last normal menstruation
postovulatory age : conceptus age
defined by the time elapsed since release of the oocyte from the ovary.
coital age : the age of a conceptus defined
by the time elapsed since the coitus that led to fertilization.
conceptional or fertilization age
: conceptus age defined by the time elapsed since fertilization. In human
clinical practice pregnancy lasts 38 weeks (266 days).
ovum : the term was formerly extended to include
any early stage of the conceptus.
Bryce-Teacher ovum : a human embryo that was thought to be the youngest
known ovum at the time of its study in 1908; now known to be a pathological
specimen.
Miller ovum : an embryo 10 or 11 days old, first described in 1913.
Peters' ovum : an embryo about 13 or 14 days old, first described
in 1899.
Mateer-Streeter ovum : an embryo about 18 days old, first described
in 1920.
Hertig-Rock ova : 34 fertilized ova, ranging from 1 to 17 days of
age, 21 of which were normal, and 13 abnormal to one degree or another;
discovered between 1938 and 1953, they constitute the only series of such
early human conceptuses in existence.
fertilization : of the 200-300 million
sperms that cross the vaginal starting line, only 250 successfully hurdle
the uterine cervix
and uterus. The hardy survivors face a grueling swim up the effectively
endless Fallopian tube thanks to thermotaxis (the upper reaches
cradling the egg are around 2°C warmer than those at the entrance).
Once in the right vicinity, sperm are thought to switch on chemotaxis,
sniffing out chemicals secreted by the egg. Without this navigation mechanism
in the female genital tract the chances of chaotic collision is negligible.
Fertilization occurs in the outer third of the Fallopian
tube.
The egg remains able to be fertilized for 24 hours after ovulation
oosperm : a recently fertilized oocyte
dispermy : the penetration of 2 sperms or spermatozoa into one oocyte.
primary zona pellucida (ZP) binding :
on egg cells : ZP glycoproteins. Sperm binding to ZP causes further activation
of cAMP/PKA and PKC, respectively. PKC opens a calcium channel in the plasma
membrane. PKA together with IP3 activate Ca2+ channels
in the outer acrosomal membrane, which leads to an increase in cytosolic
Ca2+. The depletion of Ca2+ in the acrosome will
activate a store-operated Ca2+ entry mechanism in the plasma
membrane, leading to a higher increase in cytosolic Ca2+, resulting
in F-actin dispersion which enable the outer acrosomal and the plasma membrane
to come into contact and fuse completing the acrosomal reaction.
acrosome reaction : a sequence of
structural changes that occur in spermatozoa when in the vicinity of an
oocyte in the oviduct or uterine tube, and that are believed to facilitate
entry of a spermatozoon into the oocyte: the outer membrane of the acrosome
fuses at multiple points with the overlying plasma membrane of the sperm
head, creating openings through which the hydrolases of the acrosome are
exocyted.
ADAMs (family of proteins with a disintegrin and metalloprotease domain),
including ADAM1
/ fertilin a, ADAM2
/ fertilin b and cyritestin
1 / ADAM3A, and cyritestin
2 / ADAM3B, have been investigated and found to be important for binding
rather than for fusion and painstaking studies have raised suspicions that
their putative receptors, oocyte integrins, are necessary for the sperm-oocyte
interaction.
sperm-oocyte fusion : protein phosphorylation in the flagellum.
It depends on molecules ...
Nucleoplasmin
2 (NPM2), a growing oocyte-specific nuclear protein, is crucial in
chromatin organization and post-fertilization development : analysis of
fertilization and post-fertilization events in vitro and in vivo
revealed impaired early embryo development, mostly due to failure of the
one-cell to two-cell transition, as evidenced by gross defects, including
an absence of coalesced nucleolar structures and loss of heterochromatin
and deacetylated histone H3 that normally circumscribe nucleoli in oocytes
and early zygotes, respectively.
The existence of a complex population of mRNA in human sperm is well
documented but their role is not yet elucidated. Using discontinuous density
gradients, we have isolated high and low motile sperm from the same semen
sample. The levels of different transcripts coding for molecules either
involved in nuclear condensation (protamines 1 and 2) or in capacitation
(eNOS, nNOS and c-myc) were then assessed in the 2 populations using semi-quantitative
RT-PCR. Sperm viability was estimated by eosin-nigrosin staining and by
hypo-osmotic swelling test (HOST);
apoptosis percentage was measured by the TUNEL technique. The contamination
by somatic and germ cells was assessed by looking for specific molecular
markers of these cells, respectively CD45 and E-cadherin for somatic cells
and c-kit for germ cells. The viability of sperm was unchanged in high
and low motile fractions, as well as DNA fragmentation percentage. The
amount of Prm-1 mRNA was significantly higher in low density motile than
in the high motile fraction. In most of high motile sperm samples eNOS
and nNOS transcripts were undetectable whereas they were present in the
low motile sperm. In contrast, no significant variation was found in the
c-myc/Prm-2 mRNA ratio between the two populations. Moreover, a partial
or complete disappearance of c-myc transcripts was observed after capacitation.
Thus analysing mRNA profiles could be helpful as a diagnostic tool and
prognosis value for fertilizationref.
The mRNA must exist before the
single fertilized zygote has divided into 4 and then 8 cells : after that,
you have activation of the embryonic genome, and that takes over. The egg
has its own store of mRNA, and now it has been shown that the sperm has
its own store of paternal mRNAs delivered to the egg at fertilization :
6 transcripts present in sperm, but not in unfertilized eggs, were identified
and the delivery of 2 of them—clusterin and protamine-2—into eggs was followed
using RT-PCTref.
This explains the difficulties in human parthenogenesis.
While mitochondria from the sperm are introduced into the oocyte at fertilization,
almost all of the mitochondria are excluded during the first cleavage,
and so these mRNAs may not be useful. When one made cloned animals, one
wouldn't be delivering those mRNAs. This fingerprint technology could provide
a screen for different toxicological effects to which a male may have been
exposed.
sperm aster : the centriole, with astral rays, that precedes the
male pronucleus during fertilization
fertilization membrane : a strong membrane formed around the fertilized
ovum in some species of animals by adhesion of part of the contents of
the cortical granules to the inner surface of the vitelline membrane; it
prevents the entry of additional spermatozoa.
polyspermy / polyspermia : fertilization
of 1 oocyte by > 1 spermatozoon
physiological polyspermy
: entrance of > 1 spermatozoon into an oocyte, occurring normally in certain
species, but with only one spermatozoon participating fully in the development
of the embryo
pathological polyspermy :
entrance of > 1 spermatozoon into a mature oocyte when entrance of only
one is the rule; usually development is abnormal and the embryo is not
viable. Eggs are often bombarded by innumerable sperm and all but one—the
first to fuse with the egg—must be repelled from the surface by rapidly
modifying its cell surface and extracellular matrix. Multiple variations
to this modification are found throughout phylogenyref.
The major factor blocking other male gametes from entering the zygote is
the depolarization of the membrane. That is followed by the transformation
of the vitelline layer around the embryo into a hardened fertilization
envelope (FE), which insulates the egg in a more general manner from
its environment. The driving force of this transition is secretion of the
cortical granules. At least 3 discrete steps are involved with the formation
of this extracellular barrier :
following cortical granule exocytosis, an autoactivating serine protease
severs plasma membrane attachments to the egg's vitelline layer, allowing
the separation of this matrix from the egg surfaceref
the structural components of the FE, primarily derived from the cortical
granulesref1,
ref2,
self-assemble into the vitelline layer scaffold and form a distinct pattern
of interwoven fibersref
the cortical granule-derived enzyme ovoperoxidase is targeted to the FE
by the tethering protein proteoliaisinref.
As the structural proteins self-polymerize into fibers, ovoperoxidase activity
increases in response to alkalizationref
as synthesis of one of its substrates, hydrogen peroxide (H2O2),
increasesref1,
ref2.
The catalytic activity of ovoperoxidase peaks within 5 min following fertilization,
resulting in the formation of covalent o-o-di-tyrosine bonds
between FE proteinsref1,
ref2,
ref3.
These cross-links brace the fibrous network into a solid structure, “hardening”
it to protect the embryo. An essential substrate for ovoperoxidase, and
a likely component involved with cell signaling, H2O2
is produced by a calcium-dependent mechanism involving the reduction of
one molecule of O2 and the oxidation of 2 proton donors—hereby
referred to as “oxidase activity”—whose peak output occurs at about 7–8
min postfertilizationref1,
ref2.
This temporal correlation between peak H2O2
production and ovoperoxidase activity is not coincidental; both have likely
evolved in a tightly regulated system in order to limit the potentially
lethal effects of H2O2 on the embryo without compromising
FE integrity. The existence of oxidase activity was first documented while
measuring O2 consumption following fertilizationref.
Although originally ascribed to oxidative phosphorylation—hence its misnomer
as a “respiratory burst”—the majority of the O2 consumed was
later attributed to the synthesis of H2O2. This phenomenon
was subsequently likened to the respiratory burst associated with activated
neutrophilsref1,
ref2.
Neutrophils use a complex of intramembraneous and cytosolic proteins to
regulate the oxygen reductase activity of their main player, cytochrome
b558, a heterodimeric complex of p22PHOX and gp91PHOXref.
The H2O2 synthesized by the neutrophil is used by
myeloperoxidase to synthesize the hypochlorous acid (HOCl) that kills bacteriaref.
The polymerization of this matrix is catalyzed by the enzyme ovoperoxidase,
whose indispensable substrate is peroxide. Urchin dual oxidase (Udx1),
expressed in the cell membrane of developing oocytes, is the enzyme responsible
for the typical "respiratory burst" observed at fertilization in purple
urchin (Strongylocentrotus
purpuratus) and green sea urchin (Lytechinus
variegatus). During this burst, extracellular O2 is
turned into 60 nM of H2O2, which is released
into the perivitelline space around each zygote. While other cells known
to produce H2O2—such as neutrophils, which release
peroxide to kill invading organisms—usually do not survive this synthesis,
the embryo remains immune to peroxidase. That's why the enzyme in the egg
has a dual role: at one end it's making peroxide, and at the other it's
destroying it. So it pours the peroxide outside of the cell but also doesn't
let it get inside the cell. A similar peroxidase activity can be found
in a variety of animal eggs, such as those of mice, amphibians, and mosquitoesref.
Why would a dual oxidase be favored for zygotic oxidase activity over the
multitude of other animal reductases? One key feature might be its paradoxical
and modular functions. Phylogenetic analysis of each catalytic domain shows
that the dual oxidase animal reductase domain is most closely related to
the yeast ferric reductase FRE while its peroxidase is most similar to
catalase, an enzyme responsible for the neutralization of H2O2 to H2O and
O2ref.
Our tree suggests that the enzymatic domains of the dual oxidases also
have opposing activities: calcium-dependent, pH-sensitive activation of
the animal reductase domain converts extracellular O2 to H2O2ref.
Meanwhile, the catalase-like peroxidase domain could reverse this process
to neutralize the toxic reactive oxygen species (ROS) before it can penetrate
the cell. The benefits of this antagonism can be seen by comparing the
H2O2 production of thyrocytes and eggs, both utilizing
dual oxidases, to mammalian neutrophils, utilizing gp91PHOX/NOX cytochromes
: in thyrocytes, the calcium-sensitive p138Tox/Duox2 generates H2O2
necessary for a lumenal thyroid peroxidase to conjugate iodide ions to
tyrosine residues on thyroglobulin, generating an active hormone that regulates
metabolismref.
In the sea urchin egg, Udx1 is primarily responsible for generating H2O2
necessary for ovoperoxidase-dependent cross-linking of fertilization envelope
(FE) proteins, thereby completing a competent block to polyspermyref1,
ref2,
although the H2O2 generated can be spermicidal aloneref1,
ref2
or in combination with free halides and active peroxidasesref1,
ref2.
In thyrocytes and fertilized eggs, both normally irreplaceable and nonrenewable
cell typesref1,
ref2,
the dual oxidase's catalase domain may be a first defense against H2O2
that diffuses toward the cell surface. A common second defense is cytoplasmic
enrichment in ROS scavengers such as cytoplasmic catalase, glutathione,
and glutathione peroxidase or its functional equal, ovothiolref1,
ref2,
ref3.
Neutrophils, on the other hand, directly synthesize superoxide anions (O2?)
that dismutate to H2O2 within the phagosome, where
is it used to destroy foreign invadersref1,
ref2,
ref3.
The intracellular concentration of H2O2 reaches toxic
levels, resulting in self-destruction—an acceptable outcome considering
that the pathogen is eradicated, the neutrophil is replenished by hematopoetic
stem cells, and macrophages clear the resultant debris. The insignificance
of variable cytosolic glutathione peroxidase activity on neutrophil performance
is also consistent with the dispensability of these cellsref.
Thus, the net diffusion of H2O2 away from the cell
surface by dual oxidases, versus a self-destructive phenotype, is favorable
for periodically stimulated cells that must remain viable throughout an
organism's life. Our model does not, however, limit the dual oxidase's
peroxidase domain from other activities that might make the paired enzymes
more versatile, such as protein cross-linkingref.
The maternal pool of Udx1 transcript in zygotes and the reserve pool of
Udx1 protein in granules not associated with the primary, fertilization-activated
population at the egg cortex suggest that this protein may be utilized
later in embryogenesis. One possible role for this enzyme is as a source
of cell signaling downstream of calcium. Reactive oxygen species, particularly
H2O2, have recently been identified as essential
molecules involved with autocrine and paracrine signalingref1,
ref2,
particularly in cell cycle regulationref1,
ref2
and proliferationref1,
ref2.
Given that calcium transients are necessary for passage through mitosis
and differentiationref1,
ref2,
ref3,
we postulate that the oxidase activity of Udx1 may also play an active
role in triggering H2O2-dependent signaling during
embryogensis. Regulation of oxidase activity would most likely occur through
the dual oxidase's paired EF-hand domains that are most identical to the
visinin-like protein subfamily of neuronal calcium sensing proteins, such
as neurocalcin and hippocalcinref.
Calcium sensor proteins in this calmodulin-like subfamily display free
calcium affinities in the low nanomolar to micromolar concentrations and
interact with other signaling proteins through hydrophobic residues exposed
upon calcium chelationref1,
ref2.
Therefore, multiple mechanisms would ensure that H2O2
synthesis never achieves the same flurry as the oxidative burst at fertilization:
catalase activity of Udx1, low intracellular calcium concentrationsref1,
ref2,
constitutively elevated cytosolic pHiref1,
ref2,
and regulation by PKC could all serve to modulate local concentrations
of H2O2. In turn, localized or prolonged exposure
to low concentrations of H2O2 may act to stimulate
specific cell signaling cascadesref1,
ref2,
ref3.
A survey of the heme peroxidase family identifies a cluster of ovoperoxidase-like
homologs in many invertebratesref.
In the sea urchin, this protein is expressed exclusively during oogenesisref
and requires H2O2 synthesis to cross-link the FE
constituentsref1,
ref2.
Peroxidase activity has also been identified in the oocytes and eggs of
many other species at fertilization: in mouse cortical granules and to
the outer surface of the zona pellucidaref,
in the fertilization capsule of Discoglossus pictusref,
in the chorion of the fish Tribolodon hakonensisref,
and in the eggshells of mosquitoesref.
Each of these peroxidases requires a source of H2O2.
Extrapolating from observations here, as well as the century-old history
of the “respiratory burst” at fertilization in sea urchins, we postulate
that a similar dual oxidase-dependent mechanism is functional at fertilization
throughout phylogeny.
Comparison of different peroxidase-NAD(P)H oxidase systems
(A) Unrooted phylogram of peroxidase (left) and animal reductase (right)
domains of dual oxidases compared to single-domain enzymes. Numbers are
bootstrap values for each node. Class colors correspond to specific enzymatic
modules (B–D). Bt, B. taurus; Ce, C. elegans; Ci, C. intestinalis;
Dr, D. rerio; Dm, D. melanogaster; Hr, H. roretzi;
Hs, H. sapiens; Le, L. esculentum; Lv, L. variegatus; Mm,
M.
musculus; Nb, N. benthamiana; Oc, O. cuniculus; Rn, R.
norvegicus; S. pombe, S. pombe; St, S. tuberosum;
Ss, S. scrofa. Cat/CTL, catalase; Duox, dual oxidase; EosP, eosinophil
peroxidase; FRE, ferric reductase (yeast); gp91, gp91PHOX homolog;
HRP, horseradish peroxidase; LacP, lactoperoxidase; MyeP, myeloperoxidase;
NOX, NADPH oxidase family member; OvoP, ovoperoxidase; Per, peroxidase;
Rboh, respiratory burst oxidase homolog.
Abbreviations: PKC, protein kinase C; R-Y, extracellular matrix protein
with a reactive tyrosine residue; cAMP, cyclic adenosine monophosphate;
PKA, protein kinase A; Tg-Y, thyroglobulin with a reactive tyrosine residue;
p138Tox/Duox2, thyroid NADPH oxidase/dual oxidase. H2O2,
hydrogen peroxide; NAD(P)+/H, nicotinamide adenine dinucleotide (phosphate)
in oxidized (-+) and reduced (-H) forms. For dual oxidases, note the peroxidase
domain (star) is distinct from the generic animal reductase domain (circle).
zygote : the fertilized ovum; the cell resulting
from union of a male and a female gamete (sperm and ovum). More precisely,
the cell after synapsis at the completion of fertilization until first
cleavage. Also, used loosely to refer to the fertilized ovum and early
derivatives for an indefinite period. It remains for 2-3 days in the ampulla
of Fallopian tube
and, after 3-5 days, reaches the endometrium.
nucleopetal movement : the movement of a male pronucleus toward
the female pronucleus in the zygote.
copulation path : the course taken by the male and female pronuclei
as they approach each other in a fertilized ovum.
‡ morulation => elementary, cleavage or embryonic cell / blastomere
: any one of the cells derived from the zygote by mitosis
mesentomere : a blastomere not yet divided into ...
mesomere : a blastomere of size intermediate between a macromere
and a micromere or a midzone of the mesoderm between the epimere and hypomere.
entomere : a blastomere destined to become endoderm.
cleavage / segmentation : the mitotic segmentation of the zygote,
the size of the structure remaining unchanged, as the cleavage cells, or
blastomeres, become smaller and smaller with each division.
accessory cleavage : peripheral cleavage in telolecithal oocytes
due to polyspermy.
adequal cleavage : a form in which the blastomeres are practically
equal in size.
determinate cleavage : cleavage following a precise pattern, each
blastomere having a characteristic and unalterable fate, i.e., each blastomere
becoming the precursor of a definite part of the embryo.
discoidal cleavage : cleavage limited to the animal pole of highly
telolecithal oocytes.
equal cleavage : a form in which the blastomeres are equal in size.
equatorial cleavage : cleavage that occurs in a plane passing through
the equator of the oocyte.
complete, total or holoblastic cleavage : a form in which the entire
oocyte participates in cell division
indeterminate cleavage : that following a less rigid cleavage pattern,
the blastomeres having more developmental possibilities than they usually
show, each of which, when isolated, being capable of developing into a
normal embryo.
latitudinal cleavage : cleavage in planes passing at right angles
to the oocyte axis.
meridional cleavage : cleavage in planes passing through the oocyte
axis.
incomplete, partial or meroblastic cleavage : a form in which only
the protoplasmic portions of the oocyte participate
progressive cleavage : in the formation of spores within a sporangium,
the production of a series of cleavage planes in succession, resulting
first in formation of protospores and later in formation of sporangiospores.
radial cleavage : a cleavage pattern characteristic of vertebrates
and echinoderms, in which the spindle axes are parallel or at right angles
to the polar axis of the oocyte.
spiral cleavage : a cleavage pattern characteristic of such invertebrates
as annelids and mollusks, in which the cleavage planes are oriented obliquely
to the polar axis of the oocyte.
superficial cleavage : a form in which only the surface region of
centrolecithal oocytes participate.
unequal cleavage : a form in which the blastomeres about the vegetal
pole remain larger in size than those nearer the animal pole
The first cleavage plane coincides with the plane defined by the
2 apposing pronuclei once they have moved to the centre of the egg : the
microtubule networks that allow mixing of parental chromosomes before dividing
into two may be involved in these processes. In mouse the second polar
body does not mark a stationary animal pole, but instead, in half of the
embryos, moves towards a first cleavage planeref.
The embryonic–abembryonic axis of the blastocyst arises perpendicular
to the first cleavage plane, and hence to the animal–vegetal axis of the
zygote. In most mouse embryos the progeny of one two-cell blastomere primarily
populate the embryonic part of the blastocyst and the progeny of its sister
populate the abembryonic part. However, it is not known whether the embryonic–abembryonic
axis is set up by the first cleavage itself, by polarity in the oocyte
that then sets the first cleavage plane with respect to the animal pole,
or indeed whether it can be divorced entirely from the first cleavage and
established in relation to the animal pole. The importance of the orientation
of the first cleavage was tested by imposing an elongated shape on the
zygote so that the division no longer passes close to the animal pole,
marked by the second polar body. Non-invasive lineage tracing shows that
even when the first cleavage occurs along the short axis imposed by this
experimental treatment, the progeny of the resulting two-cell blastomeres
tend to populate the respective embryonic and abembryonic parts of the
blastocyst. Thus, the first cleavage contributes to breaking the symmetry
of the embryo, generating blastomeres with different developmental characteristicsref.
morula : the solid mass of blastomeres formed
by cleavage of a zygote.
embryogenesis (see also pregnancy
for extraembryonic maternal tissues involved in development) : the production
of an embryo
embryo : in animals, those derivatives of the
zygote that will eventually become the offspring, during their period of
most rapid development, i.e., from the time the long axis appears until
all major structures are represented. In humans, the developing organism
from the fourth day after fertilization to the end of the eighth week
embryology : the science of the development
of the individual during the embryonic stage and, by extension, in several
or even all preceding and subsequent stages of the life cycle.
comparative embryology : embryology applied with a comparative view
to various species studied with reference to their taxonomy and the principle
that ontogeny recapitulates phylogeny.
descriptive embryology : the study of embryos and fetuses and their
components with reference to anatomical and chronological sequence so as
to define stages and report the course of development.
causal or experimental embryology : analysis of the factors and
relations in development, obtained by subjecting embryos to experimental
procedures
differential susceptibility : nonhomogeneity in response by the
various regions of an embryo when subjected to a diffusely applied injurious
agent
blastocyst : the mammalian conceptus in
the post-morula stage; it is like a blastula in having a fluid-filled cavity,
unlike it in having the surface layer not exclusively embryoblast but mainly
or entirely trophoblast, in having an eccentric embryoblast, and in not
being limited to one germ layer. It undergoes implantation
/ nidation
in the endometrium
inner cell mass / embryoblast
/ embryonary pole : an aggregation of cells at the embryonic pole of
the blastocyst, which is destined to form the embryo proper
formative cell : a cell of the inner cell mass of the conceptus,
a blastomere destined to form a part of the embryo
blastula /blastosphere : the usually
spherical structure produced by cleavage of a zygote, consisting of a single
layer of cells (blastoderm) surrounding a fluid-filled cavity (blastocoele)
terminal sinus : a vein which encircles the vascular area in the
blastoderm
depula : the developing egg in the stage succeeding
the blastula and preceding the gastrula.
gastrula : that early embryonic stage which
follows the blastula or blastocyst. The simplest type consists of 2 layers,
the ectoderm and the mesentoderm, and of 2 cavities, one lying between
the ectoderm and the endoderm; the other (the archenteron) formed by invagination
so as to lie within the entoderm and having an opening (the blastopore).
In human embryos the gastrula stage occurs during the third week, as the
embryonic disc becomes trilaminar.
exogastrula : a gastrula in which invagination
is hindered and the mesentoderm bulges outward.
entypy : a method of gastrulation in which
the endoderm lies external to the amniotic ectoderm.
exogastrulation : the evagination
to the exterior (or turning inside out) of the gut due to an interference
with the normal processes of gastrulation, which can occur if the morula
is cut transversely below the equator. It is usually followed by a migration
of mesenchyme cells into the interior.
mesentoderm : the inner layer of an amphibian gastrula not yet separated
into mesoderm and entoderm.
archenteron / coelenteron / gastrocoele /
primitive gut : the primordial digestive cavity of those embryonic
forms whose blastula becomes a gastrula by invagination
blastopore / archistome / protostoma / anus
of Rusconi : the opening of the archenteron to the exterior of the
embryo, at the gastrula stage
Ecker's plug : a plug of cells in the primordial mouth of the gastrula.
enteron : 1. digestive tract. 2. intestinum tenue
enterocoelic pouch : a diverticulum of the enteron of the embryo
pharyngeal septum / buccopharyngeal membrane : the transitory partition
which separates the mouth cavity from the pharynx in the embryo
tracheoesophageal septum : the septum that, during the fourth week
of embryonic development, separates the trachea from the ventral surface
of the foregut (the primordial esophagus).
Douglas' septum : the septum formed by the union of Rathke's folds,
forming the rectum of the fetus
twins : one of 2 offspring produced in the same
pregnancy
dizygotic, binovular, dissimilar, false,
fraternal, heterologous, hetero-ovular, two-egg, and unlike twins :
2 offspring developed from 2 zygotes that resulted from fertilization of
2 ova fertilized at the same time; they may be of the same or different
sex, and they have different genomes
monozygotic, enzygotic,
identical, mono-ovular, monovular, similar, true, or uniovular twins
: 2 offspring developed from 1 zygote that divided into 2 embryos at an
early stage (usually during the first 8 days); the twins therefore have
identical genomes
dichorionic twins : the separate
cells of the 2-celled embryo were to dissociate, each continuing on in
development
diamnionic or diamniotic twins : twins developing within separate
amniotic cavities
Laboratory examinations : twin peak sign
: a sonographic sign of dichorionic twinning consisting of a triangular
zone whose echotexture is similar to that of the placenta,
wider at the chorionic surface of the placenta and tapering to a point
within the intertwin membrane.
monochorionic or monochorial twins
: the inner cell mass of a normal blastocyst, which gives rise to the embryonic
axis, were to split into 2, instead of 1, masses in the blastocyst, so
that twins develop with a single chorion
monoamnionic or monoamniotic twins : twins developing within a single
amniotic cavity
diamnionic or diamniotic twins :
twins developing within separate amniotic cavities
fetal placenta / placenta foetalis / pars
fetalis placentae : the nonmaternal part of the placenta,
derived not from the fetus but from the trophoblast that envelops the fetus;
from within outward, it consists of
trophoblast : a layer of extraembryonic
ectodermal tissue on the outside of the blastocyst. It attaches the blastocyst
to the endometrium of the uterine wall and supplies nutrition to the embryo.
From it are derived the chorion and amnion.
The mesoblast, once thought to be trophoblastic, is now traced in primates
to the caudal end of the primitive streak.
cytotrophoblast / Langhans' layer
: the inner cellular layer of the trophoblast covering a chorionic villus;
it consists of ...
cytotrophoblastic cells / Langhans' cells : polygonal, mononucleate
cells resembling the cells of the cytotrophoblast, having prominent nucleoli
and clear, eosinophilic or cyanophilous cytoplasm; one of the 2 cell types
that compose a choriocarcinoma.
It secretes :
syncytiotrophoblast : the outer
syncytial layer, which consists of ...
syncytiotrophoblastic cells : large, multinucleate cells resembling
the syncytiotrophoblast, having hyperchromatic nuclei (syncytial knots
: protuberances of nuclei along the chorionic villi) and abundant eosinophilic,
sometimes vacuolated cytoplasm; one of the cell types that compose a choriocarcinoma.
It secretes
progesterone
(from maternal cholesterol, then secreted into both maternal and fetal
bloodstreams)
17b-estradiol
/ E2
and estriol
(due to placental lack of 17a-hydroxylase, they
are synthetised from androgenetic precursors in maternal circulation up
to pregnancy wk 20, then from fetal adrenal precursors (DHEA-S) processed
by placental aromatase to estrone or estradiol or by hepatic 16a-dehydroxylase
and placental sulfatase into dehydroxyepiandrosterone, then converted into
estriol by aromatase)
extraembryonic or fetal
membranes : the trophoblastic parts of the conceptus that provide for
the support of the embryo or fetus by attachment, mechanical protection,
endocrine action, and the mediation of chemical exchange with the maternal
circulation. They include :
amnion : the thin but tough extraembryonic
membrane of reptiles, birds, and mammals that lines the chorion and contains
the embryo and later the fetus, with the amniotic fluid around it; in mammals
it is derived from trophoblast by folding or splitting.
amniotic villus : one of the irregular, flat, opaque areas of imperfect
skin on the amnion near the distal end of the umbilical
cord.
amniotic sac / bag of waters : the sac
formed by the amnion
amniotic fluid / aqua amnii / liquor amnii
/ waters : fluid within the amniotic cavity produced by the amnion
during the early embryonic period, and later by the lungs and kidneys;
at first crystal clear, it later becomes cloudy. It protects the embryo
and fetus from injury; fast turnover; protective mechanical function, mobility
and constant thermal environment
amniotic fluid volume (AFV) = 500-1,500
mL in the last 10 weeks of pregnancy
in the first half of the second trimester AFV progressively increases by
10 mL/day unitl it reaches on average 500 mLat week 20, and 700 mL and
1,000 mL in the early and mid-third trimester, respectively, before decreasing
to 800-900 mL at term
817 mL at 30-week gestation (range : 318-2,100 mL)
oxalate = 19.0 (4.3) mmol/L
liquor chorii : a fluid which separates
the amnion from the chorion in the early stages of gestation.
chorion / chorial mesoblast / secundina uteri
: in human embryology, the cellular, outermost extraembryonic membrane,
composed of trophoblast lined with mesoderm; it develops villi about 2
weeks after fertilization, is vascularized by allantoic vessels a week
later, gives rise to the placenta,
and persists until birth.
chorion frondosum / shaggy or villous
chorion : the region of the chorion that bears chorionic villi
: one of the threadlike projections growing in tufts on the external surface
of the chorion
primitive or primordial chorion : the chorion from its inception
by addition of mesoderm to trophoblast through the stage in which it has
many primordial villi.
chorionic plate : the part of the inner chorionic wall in the area
of its uterine attachment, which gives rise to chorionic villi
subchorial lake or space
: the portion of the placenta, relatively free of villi, just beneath the
chorionic plate; at the edge of the placenta it becomes continuous with
irregular channels to form the marginal
lakes : discontinuous venous lacunae, relatively free of villi, near
the edge of the placenta, formed by merging of the marginal portions of
the intervillous space with the subchorial lake. Called also marginal
sinus, because it was thought to be circumferentially continuous and
important for placental drainage.
primary villus : one of the earliest chorionic villi, composed of
trophoblast only
secondary villus : an intermediate stage of chorionic villi, having
a core of connective tissue (mesoblast) covered with trophoblast
stem or tertiary villus : one of the definitive type of chorionic
villi, having trophoblastic cover, connective tissue (mesoblastic) core,
and blood vessels
branch villus : a branch of a tertiary (stem) villus through which
the main transport of substances between the mother and fetus occurs.
free or feeding villus : a chorionic villus that projects into the
intervillous
space : the space of the placenta into which the chorionic villi project
and through which maternal blood circulates
intervillous or trophoblastic lacuna : one of the blood spaces of
the placenta in which the fetal villi are found
blood lacuna : any one of the blood-filled spaces in the syncytiotrophoblast
of the embryo that serve hemotrophic nutrition.
anchoring villus / villus barbicans
: a chorionic villus that attaches to the decidua basalis
placental septum : decidual tissue
that divides the placenta into cotyledons
: any one of the subdivisions of the uterine surface of a discoidal placenta
(one of the tufted areas of a ruminant's placenta)
fetal cotyledons : 200 at end of pregnancy
maternal cotyledons : 15-20 at end of pregnancy
Within pregnancy week 10-12 cytotrophoblast cells invade the distal portion
of spiral arteries up to the border between decidua and myometrium, replacing
intimal and muscolar coats (intravascular extravillous cytotrophoblast).
Since pregnancy week 14-16 a second invasion wave replaces the intimal
and muscular coats of spiral arteries also in the intramyometrial tract,
creating unreactive, distendible, low-resistance vessels (incomplete replacement
causes preeclampsia
(PE)/eclampsia)
: uterine blood flow increases fromm 50 mL/min to 500 mL/min. After pregnancy
week 20 cytotrophoblast disappears and only syncytiotrophoblast remains.
chorion laeve / smooth chorion :
the smooth (nonvillous) and membranous part of the chorion
allantois : an initially tubular ventral
diverticulum of the hindgut of embryos of reptiles, birds, and mammals.
In reptiles and birds, it expands to a large sac for storing urine and,
after fusing with the chorion which lines the shell, provides for gas exchange.
The allantois is prominent in some mammals (carnivores, ungulates); in
others, including humans, it is vestigial except that its blood vessels
give rise to those of the umbilical cord
yolk or vitelline sac / vitellicle :
the extraembryonic membrane that connects with the midgut; at the end of
the fourth week of development it expands into the pear-shaped umbilical
vesicle connected to the body of the embryo by the long narrow yolk
stalk. In marsupial and placental mammals, it produces a complete vitelline
circulation in the early embryo and then undergoes regression; in oviparous
vertebrates, it encloses the yolk mass, breaks down yolk, and makes it
available to the developing organism. In human embryos it does not serve
a primary nutritive function, but it is the first hematopoietic
organ
of the embryo
vitellogenesis : production of vitellus / yolk : the stored
nutrient of an egg or ovum
accessory or nutritive yolk : the part of the yolk that serves for
the nutrition of the formative portion.
egg yolk : the yellow portion of the egg of a bird.
formative yolk : that part of the ovum from which the embryo is
developed, as in birds.
paraderm : the part of the vitellus of the ovum that furnishes cells
which contribute to the body of the embryo.
vitellin : a phosphoprotein found in the yolk of eggs
Eternod's sinus : a loop of vessels connecting the vessels of the
chorion with those in the underside of the yolk sac
allantochorion : a compound membrane formed by fusion of the allantois
and chorion.
chorioallantois : an extraembryonic structure derived from union
of the chorion and allantois which by means of vessels in the associated
mesoderm serves in gas exchange. In reptiles and birds, it is a membrane
apposed to the egg shell; in many mammals, it forms the placenta.
umbilical cord / funiculus umbilicalis
/ chorda umbilicalis : the flexible structure connecting the umbilicus
of the embryo and fetus with the placenta and giving passage to the umbilical
arteries and vein immersed in Wharton's jelly.
In the newborn it measures about 50-60 cm in length and has a diameter
of 15-20 mm. First formed during the fifth embryonic week from the connecting
stalk, it contains the omphalomesenteric duct (yolk stalk) and the allantois
insertion
central insertion (90%)
paramarginal insertion (<
10%)
homeobox (Hox) genes are a group of approximately 40 genes that
control the anterior-posterior axial development in all bilaterian
animals. Much interest surrounds the precise regulation of this complex
patterning, which depends inter alia upon qualitative and quantitative
differences in Hox gene product levels in different tissues at different
times. The expression pattern of this group of genes is described as temporal
and spatial colinearity as genes are expressed in time and space in
the same order that they appear on the genomeref.
Co-linearity might be achieved by 2 component steps : regulated derepression
of the locus by DNA replication, making it permissive to regulated
expression of specific transcription factors. The spatial and temporal
organization of proliferation could provide a template for developmental
gene expression in which staggered DNA replication along a developing axis
relieves gene expression from transcriptional silencing
Oct-3/4
maintains the pluripotent state of inner cell mass cells—at the blastocyst
stage of preimplantation development—that develop into the fetus after
implantation. The precise level of Oct-3/4 governs 3 distinct fates of
ES cells
a < 2-fold increase in expression causes differentiation into primitive
endoderm and mesoderm
repression of Oct-3/4 induces loss of pluripotency and dedifferentiation
to trophectoderm
A degree of Hox fragmentation has occurred in some insects, the
nematode Caenorhabditis elegans, and the tunicates Ciona intestinalis
and Oikopleura
dioica (9 locations around the genome : 8 > 250 kb from their nearest
Hox neighbor on either sideref).
Oikopleura
would resemble C. elegans, which undergoes a mosaic mode of development
where all genes are expressed early in development rather than in a temporal
sequence. Another similarity is that both have very fast development times
and a small number of cells, which might allow them to employ mechanisms
that don't require strict anterior and posterior timing. The findings also
lend support to the conjecture, first made by Darwin, that tunicates are
simplified chordates, rather than being particularly close to the chordate
ancestor. They have undergone etrograde evolution with no return : once
you have broken a cluster of genes, it's very unlikely that selection will
rebuild it. C. elegans also has a simplified body plan compared
to its ancestors.
Web resources :
primitive or primordial wandering cell : a small mononuclear cell
of the embryo that arises from the mesoderm and subsequently by differentiation
gives rise to wandering cells of the body
germinal or primitive streak :
faint white trace at the caudal end of the embryonic disc, formed by the
movement of cells at the beginning of mesoderm formation; it provides the
earliest evidence of the embryonic axis.
Hensen's protochordal or primitive node
or knot : a mass of cells at the cranial end of the primitive streak,
related to the organization of an embryo
embryonic line : the primitive streak in the center of the germinal
area.
primitive groove : a lengthwise median furrow in the primitive streak
of the embryo. In human embryos it is clearly visible during the third
week
Avian embryos have a remarkable capacity to regulate: when a pre-primitive
streak stage embryo is cut into fragments, each fragment can spontaneously
initiate formation of a complete embryonic head-tail axis, but during normal
development only one such structure typically forms. An ectopic primitive
streak can be induced by misexpression of vitellogenin
(Vg1) in the marginal zone : Vg1 induces an inhibitor that travels
across the embryo (3 mm distance) in less than 6 hours (0.5 mm per hour
: up to 10-times as fast as comparable molecules). This inhibitor acts
early in the cascade of events downstream of Vg1. FGF signalling is required
for primitive streak formation, in cooperation with Nodal
and Chordin.
3 sequential inhibitory steps ensure that a single axis develops in the
normal embryo: an early inhibitor that spreads throughout the embryo (which
can be induced by Vg1), a second inhibition by Cerberus
from the underlying hypoblast, and finally a late inhibition from Lefty
emitted by the primitive streak itselfref.
blastoneuropore : in certain embryos, a temporary aperture formed
by the coalescence of the blastopore and neuropore.
ectomesoblast : the layer of cells which has not yet become differentiated
into ectoblast and mesoblast.
mesoblast : mesoderm, especially in the
early undifferentiated stages.
segmental plate : a plate of mesoblast on either side of the notochord
at the posterior end of the embryo, from which the mesoblastic segments
are formed.
notochord / chorda dorsalis : the rod-shaped
body, composed of cells derived from the mesoblast of the primitive
node of the embryo, defining the primitive axis of the body; it is
the common factor of all species of the phylum Chordata. It is the center
of development of the axial skeleton
blastema : a group of cells that give rise
to a new individual, in asexual reproduction, or to an organ or part, in
either normal development or in regeneration.
mesoblastema / nephric blastema : all
the cells collectively which constitute the early undifferentiated mesoderm
scleroblastema : the embryonic tissue entering into the formation
of bone
germ layer theory : the theory that the embryo develops 3 primary
germ layers, each of which gives rise to definite organ derivatives.
embryonic shield : the double-layered disk from which the embryo
proper develops
germ layer : one of the 3 primary layers
of cells of the embryo (ectoderm, endoderm, or mesoderm), from which the
tissues and organs develop.
endoderm / entoderm / endoblast / entoblast :
the innermost of the 3 primary germ layers of the embryo; from it are derived
the epithelium of the pharynx,
respiratory
tract
(except the nose),
digestive
tract,
bladder,
and urethra
prechordal or prochordal plate : thickened endoderm, cephalad of
the notochord, that combines with ectoderm to become the oropharyngeal
membrane.
extraembryonic mesoderm : that located outside the embryo and belonging
to fetal accessory organs, covering for example the amnion and yolk sac.
gastral mesoderm : that infolded with the endoderm during gastrulation.
head mesoderm : loose mesoderm, cranial to the somites.
lateral mesoderm : the lateral sheets of mesoderm within which the
embryonic coelom arises.
coelom / celom / coeloma / somatic cavity :
the body cavity. In the higher invertebrates, persists throughout life.
In the mammalian embryo, it is situated between the somatopleure and the
splanchnopleure
intraembryonic coelom => principal
cavities of the trunk
extraembryonic coelom / exocoelom
: the portion of the coelom external to the embryo, bordered by chorionic
mesoderm and the mesoderm of the amnion and yolk sac; it communicates temporarily
at the umbilicus with the intraembryonic coelom
Heuser's exocoelomic membrane : a delicate sac of mesoblastic tissue
that develops as a lining of the blastocyst or chorionic cavity just after
implantation, forms the exocoelomic cavity, and quickly disappears
syncelom : the perivisceral cavities of the body considered as one
structure, including the pleural, cardiac, and peritoneal cavities, and
tunica vaginalis
paraxial mesoderm : that lying alongside the notochord
and neural tube.
peristomal mesoderm : that derived from the ventral lip of the blastopore
or from the primitive streak.
somatic mesoderm : the outer of the 2 layers into which the embryonic
mesoderm divides; associated with ectoderm to constitute the somatopleure.
splanchnic mesoderm : the inner of the 2 layers into which the embryonic
mesoderm divides; associated with endoderm to constitute splanchnopleure.
somatopleure : the embryonic body wall, formed by ectoderm and somatic
mesoderm.
splanchnopleure / splanchnoderm : the layer formed by the union
of the splanchnic mesoderm with endoderm; from it are developed the muscles
and the connective tissue of the digestive
tract
somite / mesoblastic, mesodermic, primitive, primordial,
or protovertebral segment : one of the paired, blocklike masses of
mesoderm, arranged segmentally alongside the neural
tube of the embryo, forming the vertebral
column
and segmental musculature
dermomyotome : all but the sclerotome
of a mesodermal somite; the primordium of skeletal muscle and, perhaps,
of corium.
dermomyotome plate : the portion of the embryonic somite remaining
after migration of the sclerotomic tissue
myotome / myomere : the muscle plate
or portion of a somite that develops into striated (skeletal) muscle. A
group of muscles innervated from a single spinal segment.
hypomere : the ventrolateral portion of a myotome, innervated by
an anterior ramus of a spinal nerve. The lateral plate of mesoderm that
develops into the walls of the body cavities.
epimere : the dorsal portion of a somite, from which is formed muscles
innervated by the dorsal ramus of a spinal nerve.
sclerotome : one of the paired masses of
mesenchymal tissue, separated from the ventromedial part of a somite, which
develop into vertebrae and ribs
ethmovomerine plate : the central part of the ethmoid bone in the
fetus.
foot plate : a flattened expansion at the end of the limb end of
the embryo, the precursor of the foot
frontal plate : a fetal plate of cartilage between the sides of
the ethmoid cartilage and the sphenoid bone.
frontonasal plate : a fetal plate from which the external nose is
developed
hand plate : a flattened expansion at the end of the limb bud of
the embryo, the precursor of the hand
lateral mesoblastic plate : the thickened portion of either side
of the mesoblast.
subgerminal plate : a sheet of protoplasm forming the floor of the
segmentation cavity of the ovum.
urethral plate : an endodermal plate that gives rise to the terminal
portion of the spongy urethra
epiblast : the upper layer of the bilaminar embryonic disc present
during the second week; it gives rise to ectoderm; ectoderm; the ectoderm
except for the neural plate.
ectental line : the line of junction between the ectoderm and endoderm
ectoderm / ectoblast : the outermost layer
of cells of the 3 primary germ layers of the embryo; an outer protective
coat. From it are developed the epidermis
and the epidermal tissues, such as the nails,
hair,
enamel
of teeth,
and glands
of the skin,
the nervous system,
the external sense organs such as the ear
and eye,
and the mucous membrane of the mouth
and anus
amniotic ectoderm : the inner layer of the amnion (and covering
of the umbilical cord) that is continuous
with body ectoderm.
Wharton's jelly : the soft, jellylike,
homogeneous intercellular substance of the umbilical
cord; it gives the reaction for mucin and contains thin collagenous
fibers which increase in number with the age of the fetus.
basal ectoderm : trophoblast covering the eroded uterine tissue
that faces the placental sinuses.
blastodermic, primitive or primordial ectoderm : the external layer
of a blastula or blastodisk
chorionic ectoderm : the trophoblast.
extraembryonic ectoderm : a derivative of epiblast or ectoderm located
outside the body of the embryo.
dermatomere : any segment or metamere of the embryonic integument.
dorsal flexure : one of the flexures of the embryo in the mid-dorsal
region
caudal or sacral flexure : the bend at the aboral (caudal) end of
the embryo
Rathke's craniobuccal, craniopharyngeal
or hypophyseal pouch or pocket : a diverticulum from the embryonic
buccal cavity, from which the adenohypophysis is developed; its lumen persists
in adults as small colloid-filled cysts and clefts at the juncture of the
pars distalis and the neurohypophysis
bucconasal or oronasal
membrane : a thin epithelial plate separating the nasal pits from the
oral cavity of the embryo
Seessel's pouch : a transient outpouching
of the embryonic pharynx rostrad of the oropharyngeal membrane and caudal
to Rathke's pouch.
branchial or visceral arches
: paired arched columns that bear the gills in lower aquatic vertebrates
and that, in the embryos of higher vertebrates, appear in comparable form
before subsequent modification into structures of the head and neck. In
humans they are also called pharyngeal arches because gills do not
develop. Each contains a cartilaginous bar, consisting of right and left
halves.
the first pharyngeal arch (mandibular
arch) is innervated by the V cranial nerve and vascularized by the
external maxillary artery
the dorsal surface differentiates into the body of stapes, the head of
malleus, and the ear pavillon
the ventral surface differentiates into the Meckel's cartilage and the
malleus
the median line of the pharynx differentates into the body of tongue
sphenomandibular and anterior malleolar ligaments, malleus, and incus ?
the second pharyngeal arch (hyoid
arch) is innervated by the VII and VIII cranial nerves and is vascularized
by the stapedial artery
the dorsal surface differentitates into the stapes
the ventral surface differentiates into the stylohyoid ligament, the styloid
process and the lesser horn of the hyoid bone, and cranial part of the
hyoid body
the median line of the pharynx differentates into the root of the tongue,
the foramen caecum, and the median part of the thyroid
the third pharyngeal arch is innervated
by the IX cranial nerve and vascularized by the carotid artery. The ventral
surface differentiates into the greater horn of the hyoid bone and the
caudal part of its body
the fourth pharyngeal arch is innervated
by the X cranial nerve and vascularized by the aortic arch at left and
part of the subclavian artery at right. Its ventral surface differentiates
into the thyroid, cuneiform and part of the epiglottic cartilages
the sixth pharyngeal arch differentiates into the lateral regions
of the thyroid
the actual sixth pharyngeal arch is innervated by the inferior laryngeal
nerve and vascularized by the pulmonary artery and the Botallo's ductus
arteriosus. It differentiates into the cricoid, aritenoid and corniculate
cartilages. In the human embryo, the sixth arch is actually the fifth in
number but is so named for reasons of comparative anatomy and evolution;
it does not appear on the surface
The second branchial arch develops far more than the caudal ones thanks
to the nuchal curvature and covers the third and forth at the end of week
5. Sinus precervicalis : the depression at the side of the neck,
produced in the developing embryo by the growth of the pharyngeal (branchial)
arches.
branchial or pharyngeal cleft, fissure
or groove : the embryonic ectodermal cleft between successive pharyngeal
arches
first cleft differentiates into the external auditory meatus
The second branchial cleft deepens and forms the cervical sinus
(a temporary depression caudal to the embryonic hyoid arch, containing
the succeeding pharyngeal arches), which closes off as the cervical
vesicle.
branchial, pharyngeal or visceral pouch
: a lateral diverticulum of the pharynx that meets a corresponding pharyngeal
groove in the embryonic ectoderm, forming a closing membrane that may rupture
and complete the gill slit as observed in lower vertebrates
first pouch differentiates into the Eustachian tube, the cavity
of middle ear, and the mastoid cells
second pouch differentiates into the palatine tonsils and the supratonsillary
fossa
third pouch
the dorsal surface differentiates into the lower parathyroids and the pyriform
fossa
the ventral surface differentiates into the thymus
forth pouch
fifth pouch differentiates into the lateral regions of the thyroid
ultimobranchial, postbranchial
or telobranchial bodies : embryonic derivatives of the fifth pharyngeal
pouches, which migrate along with the parathyroid
glands
and are incorporated in the thyroid
gland.
In submammalian vertebrates they remain as discrete masses in the neck
or mediastinum throughout adult life. The parafollicular
cells
of these bodies produce calcitonin.
branchial sinus : an abnormal opening between a pharyngeal groove
and its corresponding pharyngeal pouch, homologous with an ancestral gill
slit
branchial cleft : any of the slitlike
openings in the gills of fishes, formed between the branchial arches
hyobranchial or posthyoidean cleft : the cleft between the hyoid
and the next succeeding arch in the developing embryo
hyomandibular or hyoid cleft : the cleft between the mandibular
and hyoid arches in the developing embryo
epithelial plug : a mass of ectodermal cells that temporarily closes
the external naris of the fetus.
neural arch : one of the cartilaginous structures surrounding the
embryonic spinal cord, formed by the dorsad growth of the dorsolateral
arcualia; it is the primordium of the vertebral arch
hemal arch : one of the cartilaginous structures
surrounding the caudal vein in the tail of the vertebrate embryo, formed
by the ventrad growth of the ventrolateral arcualia. In fish, the arches
are also present in the thoracic region
pleuropericardial membrane
: a membrane in the embryo separating the pericardial cavity from the pleural
cavity
pleuroperitoneal membrane
: a membrane in the embryo separating the pleural cavity from the peritoneal
cavity and developing into the posterolateral part of the diaphragmù
neuroepithelium / neurepithelium / sensory
epithelium : simple columnar epithelium made up of cells specialized
to serve as sensory cells for the reception of external stimuli, as the
sensory cells of the
neuroepithelium macularum / neuroepithelium of maculae : the specialized
epithelium of the maculae of the labyrinth, containing receptor cells from
some of which cilia (sensory hairs) project into the statoconic membrane.
neuroepithelium cristae ampullaris / neuroepithelium of ampullary crest
: the specialized epithelium of the ampullary crest of the labyrinth, containing
receptor cells from some of which cilia (sensory hairs) project into the
cupula.
The epithelium of the ectoderm, from which the CNS is developed.
at week
4, epiblasts migrated into hypoblast differentiate into primordial germ
cells (PGCs) sized 15-20 mm : germ cell
fate in mice is induced in pluripotent epiblast cells in response to signals
from extraembryonic tissues. The specification of approximately 40 founder
PGCs and their segregation from somatic neighbours are important events
in early development. A critical event during this specification includes
repression of a somatic programme that is adopted by neighbouring cells.
Blimp1 (also known as Prdm1), a known transcriptional repressor, has a
critical role in the foundation of the mouse germ cell lineage, as its
disruption causes a block early in the process of PGC formation. Blimp1-/-
mutant embryos form a tight cluster of about 20 PGC-like cells, which fail
to show the characteristic migration, proliferation and consistent repression
of homeobox genes that normally accompany specification of PGCs. Furthermore,
our genetic lineage-tracing experiments indicate that the Blimp1+
cells originating from the proximal posterior epiblast cells are indeed
the lineage-restricted primordial germ cell precursorsref.
At week 5 they begin migrate from the splanchnopleure of the secondary
yolk sac through the mesenterium with ameboid movements into the genital
ridge. The compartmentalisation of the PGCsis completed in
humans by the 7th week of gestation
ventrolateral mass : that portion of the primordial lateral mass
of the embryo from which are developed the abdominal, thoracic, and anterior
cervical muscles.
bud : any small part of the embryo or adult metazoon more or less
resembling the bud of a plant and presumed to have potential for growth
and differentiation.
bronchial bud : an outgrowth from the stem bronchus giving rise
to the air passages of its respective pulmonary lobe.
end or tail bud : the remnant of the primitive knot, from which
arises the caudal portion of the trunk
limb bud : a swelling on the trunk of the embryo that becomes a
limb.
lung bud : an outgrowth from the foregut that gives rise to the
trachea, bronchi, and all the branchings that form a tracheobronchial tree.
periosteal bud : vascular connective tissue from the periosteum
growing through apertures in the periosteal bone collar into the cartilaginous
matrix of the primary center of ossification.
tail bud : the primordium of the caudal appendage
metanephric or ureteric bud or diverticulum : an outgrowth of the
mesonephric duct that gives rise to all but the nephrons of the permanent
kidney
at weeks 5 the cloacal fold divides into anal membrane and genital fold
sexual differentiation :
determination of chromosomal or genetic sex at fertilization
formation of organs common to both sexes
sacrogenital fold / plica recto-uterina
/ plica rectouterina [Douglasi] / rectouterine fold : a crescentic
fold of peritoneum extending from the rectum to the base of the broad ligament
on either side, forming the rectouterine pouch
urogenital fold or ridge : a longitudinal
ridge or fold in the embryo, lateral to the root of the dorsal mesentery,
which later subdivides longitudinally into ...
wolffian or mesonephric ridge : the more lateral portion of the
urogenital ridge, which gives rise to the mesonephros.
rete, medullary or sex cords : strands
of primordial cells in the medulla of the embryonic gonads that connect
with some of the mesonephric tubules, and from which the rete
ovarii
or the rete testis
develops
genital or germ ridge or fold (bipotential
gonads) : the more medial portion of the urogenital ridge, which gives
rise to determination of gonadal sex
ductus paramesonephricus / gasserian, müllerian,
paramesonephric or primordial duct / ductus muelleri / duct of Müller
(either of the paired embryonic ducts arising as a peritoneal pocket, extending
caudally to join the urogenital sinus) are created by default
external genitalia arise as feminine
urogenital sinus / sinus urogenitalis
: an elongated sac formed by division of the cloaca in the early embryo,
communicating with the mesonephric ducts and bladder, and forming the vestibule
of vagina,
urethra
and lower vagina
in the female and prostate
and posterior urethra
in the male
genital tubercle : an eminence ventral
to the cloaca in the early embryo, which is the primordium of the glans
penis
or the clitoris.
genital or labioscrotal
swelling : an elevation on each side of the embryonic phallus that
becomes either a labial (labia majora) or a scrotal swelling
determination of internal ducts => determination of phenotypic sex
in embryos with no Y chromosome (ie genetic females)
interstitial Leydig cells are stimulated by LH
or hCG
to produce testosterone,
which induces
homolateral formation of ductus mesonephricus
/ mesonephric duct / ductus Wolffi / wolffian duct / duct of Wolff / Leydig's
duct / canal of Oken (an embryonic duct which, initiated in association
with rudiments of the pronephros,
is taken over as an excretory duct by the mesonephros
and metanephros);
later it differentiates into :
urethral groove : the embryonic groove that becomes the penile urethra
as the urogenital folds on each side fuse.
Testicles might be outside the body because temperature influences the
sex of human children. A temperature-sensitive gender switch that makes
hot sperm male could be a relic from our evolutionary past, given the evidence
that more males are born in hot climates. Temperature determines sex in
many reptiles. For example, alligators that develop at or below 30ºC
turn out to be female, whereas hotter eggs yield males. Such a system wouldn't
work in mammals: they maintain a constant body temperature and carry their
young within. Instead, a gene called SRY on the male-only Y chromosome
kicks off male development in embryos. This genetic switch might have been
bolted on to the front of a redundant, but still functioning, temperature-driven
mechanism. Male animals are sometimes born without functioning SRY, suggesting
that something else - perhaps temperature - might be able to trigger male
development. High temperatures make males by activating genes that speed
up embryonic development : males are known to develop more quickly than
females. It's usually thought that testicles are kept away from the body's
hot core because sperm grow best in cool conditions. This makes little
sense as inventing the ability to survive at a slightly higher temperature
seems easier than having external testicles : males' cool gonads are painfully
vulnerable, their descent is a dicey process that can lead to hernias,
and the testicles' position might instead prevent overheated sperm going
down the road to maleness. Sperm's low tolerance for heat, which develops
only after birth, might be an additional safeguard to keep the sex switch
inactive. There are many possible explanations for the paradoxes of maleness
besides temperature : for example, it might be just as easy to move the
temperature switch to a hotter point as to avoid it by moving testicles
outside the body. But the idea that the sex switch is still there is testable
by keeping sperm in tissue culture at different temperatures, in the hope
of spotting the narrow window between activating a maleness switch and
killing the sperm. The human Y chromosome contains 60 Mbp of DNA, it is
haploid, and 95% of it is nonrecombining : the 23Mb euchromatic male-specific
region (MSR) in the Y chromosome comprises 8 massive palindromic
sequences and these regions are rich in genes that are functional and
testis-specific : they contain at least 156 transcription units, all located
within euchromatic sequences, and 24 MSY-specific families account for
125 of these. Half of the transcription units encode 27 distinct proteins
or protein families, 12 of which are expressed ubiquitously, and 11 of
which are testis-specific, confirming a previous model proposing 2 distinct
functional classes of