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Developmental Biology
Kathi McDowell, NSUBA, Spring 2013
120
Biology
Undergraduate 4
01/15/2013

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Term
Hans Spemann
Definition
German who tied hair loop between developing tadpole cells and showed that not always equivalent cells
Term
second messenger
Definition
activation of a relay by a receptor on a cell receiving an initial cellular signal
Term
signal transduction
Definition
the process in which a signal evokes a cellular response
Term
ligand
Definition
that which binds to or is tied to a receptor
Term
endocrine
Definition
ligand is secreted and enters the circulation
Term
paracrine
Definition
ligand is secreted but remains in the locality to affect nearby cells
Term
autocrine
Definition
ligand secreted to act back on the signaling cell itself
Term
Lineage
Definition
traces lines of descent. Unequal division of cytoplasmic material can explain differences between related cells
Term
Position
Definition
Examines the neighboring cell populations. Autocrine and paracrine signaling can influence cells. Variations in temperature, UV light, ion concentration, etc.
Term
morphogens
Definition
maternal effect factors derived from mother's genetic information - transcribed or translated and then stored in eggs
Term
totipotent
Definition
entire embryo and fetal placenta
Term
pluripotent
Definition
all the embryonic structure
Term
John Gurdon
Definition
Nuclear equivlanence
Term
Oogenesis
Definition
oogonium - primary oocyte - secondary oocyte - ootid - ovum
Term
spermatogenesis
Definition
spermatoonium - primary spermatocyte - secondary spermatocyte - spermatids - spermatazoon
Term
capacitation
Definition
female secretions that make sperm viable
Term
marine sperm
Definition
acrosomal filament - protein binding - species specific
Term
syngamy
Definition
still have 2 separate pronuclei in egg
Term
leydig cells
Definition
support cells that provide sperm testosterone
Term
seritoly cells
Definition
support cells that provide nutrients to sperm
Term
primordial germ cells
Definition
give rise to sperm and eggs, migrate to gonads, not originally formed there.
Term
germinal vesicle
Definition
the developing primary oocyte enters the extended prophase of the first meiotic division, it is exceptionally large, with a very prominent nucleus often called the germinal vesicle
Term
Pronucleus
Definition
The nucleus of the egg from the two divisions of meiosis is called female pronucleus - it contains only one of each chromosome type and is haploid.
Term
Seminiferous tubules
Definition
Germ cells and their associated accessory cells line the perimeter of connective tissue tubes called seminiferous tubules.
Term
spermiogenesis
- histones replaced by protamines (dense packing of chromatin)
Definition
spermatids undergo reorganization and transition from spermatid to spermatozoon near the center of the seminiferous tubule
Term
Nuclear equivalence
Definition
Take any cell and put it into an activated egg. It will produce a new cloned organism.

Genomic (nuclear) equivalence – Each cell has the same complement of DNA

Nuclei from differentiated adult cells are able to give rise to all embryonic tissues

This new embryo can become an adult – usually stays as tadpole

Gurden, et al.
1-Culture frog skin cells
2-Break membrane by drawing the cells into a narrow pipette
3-Take an activated frog egg and functionally remove the nucleus, i.e. UV light to inactivate nucleus.
4-Inject cells into enucleated egg
5-Cleavage occurs
6-Tadpole forms - shows nuclei are totipotent
7-Rarely get adult - something else happening. Also done with insects & Dolly the famous sheep
Term
Totipotent stem cells
Definition
Each cell has the ability to form the entire embryo plus fetal placenta

Totipotent stem cells can form embryo and fetal placenta
Term
Pluripotent stem cells
Definition
form the embryo

Pluripotent stem cells – Each cell has the ability to form all the embryonic structure
Term
Committed stem cells
Definition
Can give rise to subsets of cell types

Committed stem cells can give rise to subsets of cell types, i.e. hemangioblasts - blood vessels, blood cells and lymphocytes
Term
Progenitor cells
Definition
are precursor cells. They have some level of differentiation. They do not divide into progenitor cells, i.e. Myeloid progenitor cells - can generate all different type of blood cells.

Progenitor (precursor) – no longer stem cells – give rise to very specific types of cells yet can still be considered general
Term
Nonequivalent cell division
Definition
Mother cell can change mating type
Daughter cell cannot
Divide a Frog egg at 2 cell stage. Sometimes get two embryos. Sometimes get one embryo and one poorly differentiated entity.

Mouse Embryo
Outer cells can form extra-embryonic tissue
Inner cells form embryo

16 cell stage mouse embryo
A Cells - placenta - therefore committed cells
B Cells - embryo - therefore pluripotent cells

These A and B cells are nonequivalent
Term
Lineage
Definition
traces lines of descent. Unequal division of cytoplasmic material can explain differences between related cells

Lineage traces lines of descent  cytoplasm and subcellular organelles shared by offspring. Not necessarily equivalent.
Term
Position
Definition
Examines the neighboring cell populations. Autocrine and paracrine signaling can influence cells. Variations in temperature, UV light, ion concentration, etc.

Position Information obtained from neighboring cells or environment in which cell is located.
Term
Regeneration
Definition
Lose a body part
Neighboring cells dedifferentiate
area expands and cells redifferentiate

Blastema forms

The nuclei of cells are equivalent yet cells have specific fates depending on environment.

During regeneration the cells dedifferentiate and start to follow a different path.

Sever a limb from a salamander, the stump heals and a mound of cells called the blastema form under the epithelium and over the stump. Get cell division, migration, redifferentiation and a new intact limb.

Note: Differentiation is when cells become specialized. Some genes are active and others inactive in a specific cell type.
Term
Environmental Influences
Definition
Environmental influences internal Vs external, i.e. signaling from outside, influences internal environment. Maternal effect factors (distribution of factors) influences different regions of the embryo.

As we will find out later even the site of sperm penetration causes environmental changes.
Term
Zygote
Definition
single fertilized egg
Term
Metamorphosis
Definition
Larvae to adult
Term
Gonads
Definition
Organs for reproduction
Ovary – egg formation

Testis – sperm formation

Primordial germ cells (PGC’s)
Term
Gametogenesis
Definition
Formation of gametes (sperm and egg). The gametes are formed in the gonads. The gonads are the ovaries and the testes. An ovary produces eggs and a testis produces sperm

Primordial germ cells (PGC’s) are the precursor to the germ line. PGC’s do not arise in gonads but migrate to developing gonads.

Most animals have a clear separation of germ cells and somatic cells early in development. This helps to keep the germ line safe.

For other animals and plants this is not the case. For example in jellyfish (cnidaria), sea squirts (tunicates) and flat worms the somatic cells can become germ cells even in adults.
Term
Oogenesis
Definition
Oogonia
PGC’s within ovary
Stem cells in frogs
Not stem cells in humans
Accessory cells
Contribute to egg development
Support cells
Produce hormones
Produce shells
Release egg from ovary

Extensive growth
Germinal vesicle formation
Meiosis
Equal karyokinesis
Unequal cytokinesis

Oogenesis is characterized by extensive growth, i.e. chicken, reptile – single cell but lots of yolk.

Germinal vesicle formation. Some organisms form a germinal vesicle. This is when the developing egg stops after it undergoes Meiosis I. Frog oocytes have germinal vesicles.

Pronucleus formed. Other organisms form a pronucleus. This occurs when the 2nd Meiotic division is completed. Humans form a pronucleus
Term
The first step in gametogenesis
Definition
get PGC to genital ridge as gonad is forming (both male and female)

PGC in ovary called oogonia In humans these are not stem cells. A single oogonium will mature into a Primary oocyte. In Frogs these are stem cells. Each season the oogonium will divide via mitosis into 1000’s of Primary oocytes

Accessory cells - contribute to oogonium .

Support cells produce hormones, make shells, and release the egg from the ovary.
Term
Granulosa
Definition
Accessory cells - contribute to oogonium
Term
theca
Definition
Supporting cells - produce hormones, make shells, and release the egg from the ovary
Term
Germinal vesicle formation
Definition
Some organisms form a germinal vesicle. This is when the developing egg stops after it undergoes Meiosis I. Frog oocytes have germinal vesicles.

Xenopus laevis
Term
Pronucleus formed
Definition
Other organisms form a pronucleus. This occurs when the 2nd Meiotic division is completed. Humans form a pronucleus
Term
Highly Organized Egg
Definition
Stores vast quantities of material
Yolk assembled into organelles called yolk platelets – unevenly distributed to vegetal side (animal side less yolk)

Pigment granules, if present, may or may not be evenly distributed. The frog has pigment granules in its animal hemisphere to protect from UV light.

Female pronucleus (or germinal vesicle) is usually near the periphery.

Cortical granules near surface.

Cortex - Peripheral cytoplasm – has different cytoskelatal structure than internal cytoplasm.

NOTE: Parts of egg very differently, therefore future mitotic cells will have different environments.
Term
Spermatogenesis
Definition
is the production of sperm from Primordial Germ Cells.

PGC’s arrive at genital ridge
Incorporate into sex chords
At maturity-sex chords hollow – seminiferous tubules
Epithelium differentiate into Serotoli cells
PGC’s become type A1 spermatogonium
Leydig cells - testosterone

In male vertebrate
PGC’s arrive at genital ridge. They became incorporated into sex chords and stay there until maturity. At maturity the sex chords hollow to form seminiferous tubules. The epithelium of the seminiferous tubules differentiate into sertoli cells (accessory cells). These cells nourish and protect developing sperm cells.

PGC’s become type A1 spermatogonium (stem cells). These are smaller than PGC’s and are characterized by ovoid nucleus.
A1 Spermatogonium are found adjacent to outer membrane of sex chords.

During fetal development Interstitial Mesenchyme cells of testes differentiate into leydig cells (support cells) which produce testosterone

A1 spermatogonium divide-Produce A1 (stem cells) and A2 spermatogonium
A2 spermatogonium divide producing 2x A3 spermatogonium
A3 spermatogonium divide – each producing 2x A4 spermatogonium (still stem cells)

A4 spermatogonium divide
New A4 spermatogonium
apoptosis
Intermediate spermatogonium (committed stem cells)
Undergoes mitosis – 2x Type B spermatogonium
Undergoes mitosis – 2x Primary spermatocyte
Undergoes meiosis yielding spermatid

A4 - New A4 Spermatogonium
Or Apoptosis
Or Intermediate Spermatogonium

Intermediate Spermatogonium – Committed Stem Cell
- Type B Spermatogonium  1 Spermatocyte
- Type B Spermatogonium  1 Spermatocyte

1 Spermatocyte  Undergoes Meiosis (yielding Spermatids)
Term
Fate Maps and Lineage Diagrams
Definition
Trace cell lineage
Cell – Cell interaction

Lineage and position play a role in determination.

Fate maps are prepared by tagging cells and then observing the location of the descendents in the adult. These can be displayed as a lineage map which outlines the descendents of the original cells and their final structure within the adult
Term
Cleavage
Definition
rapid mitotic division after fertilization
Term
Holoblastic
Definition
isolecithal (limited or no yolk) and mesolecithal (moderate amount of yolk) eggs

Entire cytoplasm of Zygote is partitioned into cells. Seen in species with low to moderate yolk content
Term
Radial cleavage
Definition
Echinoderms and amphibians (mesolecithal - displaced radial)
Term
Spiral cleavage
Definition
Annelids, mulluscs, and flatworms
Term
Bilateral cleavage
Definition
Tunicates
Term
Rotational cleavage
Definition
Mammals and nematodes
Term
Morphogens
Definition
are maternal effect genes which produce maternal effect gene products. By definition - Morphogens – substances (proteins) that in influence development by their presence, absence, and in between concentration (gradients)

There are 34 Morphogens in Drosophila
12 Anterior – Posterior (9 of these are Posterior)
6 Extreme ends, Acron & Telson
16 Dorsoventral
Term
Convergent Extension
Definition
Simultaneous narrowing and lengthening of a sheet of cells. Brought about by intercalation of cells.
Term
Ingression
Definition
Migration of individual cells from surface layer to the interior of embryo.
Term
Invagination
Definition
Movement of a sheet of cells into the interior of the embryo, forming a pocket which has an opening to the surface.
Term
Involution
Definition
Folding of a sheet of cells under itself. The sheet then spreads over the internal surface of the cells that remain on the outside.
Term
Epiboly
Definition
Spreading of a sheet of cells to enclose underlying layers. It is brought about by increase in cell numbers (cell division) flattening of cells or intercalation of subsurface cells into the surface layer
Term
Delamination
Definition
Splitting of a layer of cells into 2 layers
Term
Four extraembryonic membranes
Definition
Chorion – outer layer of protection
Allantois – Highly vascularized
Yolk sac – highly vascularized
Amnion –inner layer of protestion
Term
Spemann Organizer (Dorsal lip of Blastopore)
Definition
A) In older view - induces the nervous system

B) In current view - inhibits cells from becoming epidermis. Therefore the default structure is neural tissue.

Ventral portion of the embryo’s ectoderm produces BMP4 (BMP family of growth factors).

Dorsal ectoderm releases Noggin and Chordin. Chordin is an antagonist to BMP4. Noggin and Chordin form concentration gradient in gastrula phase. Therefore the organizer tissue in the dorsal midline have higher anti BMP concentration and lower BMP concentration. This prevents prospective neural plate cells from becoming epidermis. They form neural tissue by default.

Not sure what causes differentiation between brain and spinal chord. We will discuss this more later in the semester.
Term
SAM
Definition
Shoot apical meristem
Term
AB
Definition
Auxillary bud
Term
Cotyledons
Definition
Seed leaves
Term
Epicotyl
Definition
Stem above the cotyledons
Term
Hypocotyl
Definition
Stem portion between cotyledon and root
Term
Leaf Axil
Definition
Junction between main stem and leaf
Term
Meristems
Definition
regions of cell division. They are embryonic like throughout the life of plant. The meristems are constantly generating new plant organs.
Term
Plants cell fate - position vs. liveage
Definition
In plants position not lineage determines cell fate
Term
Apical Meristem
Definition
Growing Tip of Shoot and Root

The center is the Central Zone and consists of cells with lots of small vacuoles that renew the meristem – slow cell division.

Peripheral Zone – surrounds the Central Zone – is the morphogenetic region. It contains cells that form the organs – rapid cell division.
Term
Outer Layers L1 and L2
Definition
Tunica
Term
Interior Layers L3
Definition
Corpus
Term
Chimeric plants can be used to study final destination of cells.
Definition
The destinations are as follows:
L1 – Epidermis

L2 – Bulk of leaf and flower

L3 – Central portion of organ and stem
Term
Phyllotaxy
Definition
position of leave on the stem. Involves communication among existing and newly formed leaf primordium.

Phyllotaxy may change as the plant matures (i.e. spiral  Whorled) or by external manipulation (cut the meristems)
Term
Leaves arranged as spiral or whorls
Definition
Whorls can have one leaf, 2 leaves or 3 leaves (leaves/node). The nodes are alternating (180). This is due to chemical and physical interaction and constraints. For instance maize is normally 1 leaf alternating but the mutation abphyl (aberrant phyllotaxy), gives a very large meristem with 2 leaves instead of one.
Term
Auxins
Definition
allow for cell elongation or expansion. This causes cell wall expansion leading to change in organ pattern.

Class of hormones.
Term
Axillary Meristems
Definition
Leaf axil is the junction between leaf & main stem. Axillary meristems are found in leaf axils and produce axillary buds.

SAM displays apical dominance – suppress axillary meristems get Christmas tree appearance.

Remove SAM get bushy appearance
Term
Adventitious meristems
Definition
forms from surface cells on stems and leaves get bushy appearance even without SAM
Term
Cylinder type meristems
Definition
cambrium layer causes increase in width of plants or production of bark.
Term
Root cap
Definition
is for protection
Term
Zone of Cell Division
Definition
is the meristematic region just below the root cap
Term
Zone of Cell Elongation
Definition
is where the cells elongate
Term
Zone of Cell Differentiation
Definition
is where cells differentiate into Xylem and Phloem as well as where lateral roots and root hairs form
Term
Quiescent Center of Root Meristem
Definition
contains slowly dividing cells
Term
Reproduction
Definition
Ancestral angiosperms are believed to have formed a terminal flower directly
from SAM. Modern angiosperm – variety of flowing patterns where axillary buds
are inflorescent – Inflorescent meristem shift from making leaves to making
flowers. The change to making flowers probably arises from centroradiulus
(cen) gene (in snapdragon). CEN prevents terminal flower formation – CEN
suppresses Floricaula (Flo) (specifies floral meristem) however, FLO stimulates CEN
Term
photoperiod
Definition
Some flowering is regulated by day length
Term
Where two cortex cells meet
Definition
will be root hair (Epidermis forms root hairs)
or atrichoblasts

Root hairs absorb water, allow bacteria to enter, grows in zone of differentiation from epidermal cells.
Term
Glabra 2 (gl2) mutant
Definition
– all epidermal cells make root hairs
Hairy roots, no hair on stems.
Term
Normally atrichoblasts produce gl2
Definition
which produce atrichoblasts
Term
Hair cell precusors form
Definition
Trichoblasts (hair precursors) and are structurally different than atrichoblasts (non-hair epidermal cells).

Cortical cells provide signal. Hair cells are next to 2 cortical cells whereas non-hair cells are next to one.
Term
Trichomes
Definition
Hair on leaves & stems

gl2 mutants produce trichomes on leaves which are small compared to normal
and the plant has hairy roots.
Term
Transparent testa glubra (ttg) mutant

Why?
Definition
no trichomes on leaves, lots of hair on roots.

ttg stimulates gl2. No ttg then no gl2
Term
Werewolf (wer)
Definition
expressed in non hair cells (root) are required for differentiation.
Term
Glabral 1 (gl1)
Definition
is expressed in the epidermal cells of leaf and are required for trichome formation.
Term
In root, wer, and in leaf, gl1
Definition
are needed to stimulate gl2
Term
Asymmetrical division yields
Stomata Formation
Definition
guard mother cell (GMC)

Adjacent cells undergo asymmetric division, smaller ones close to GMC called subsidiary cells
GMC divides symmetrically producing guard cells

Stomata formation occurs in both monocots and dicots, The stomata is an opening surrounded by photosynthetic guard cells. When the guard cells are actively photosynthesizing (daytime) the stomata is open. It is closed at night.
Term
Plant Hormones
Definition
Abscisic acid
Auxin
Brassinosteroids
Cytokinins
Ethylene
Giberillins
Term
Abscisic Acid (ABA)
Definition
Stomata closure, maintenance of dormancy. Prevents germination. Soak seeds leach out ABA get germination.
Term
Auxin
Definition
Apical dominance, cell elongation, phototropism, geotropism, fruit development. Xylem regeneration in wound healing, adventitous root formation.
Term
Brassinosteroids
Definition
Cell elongation, cell division. Produces steroid reductase enzyme (in humans this enzyme would reduce testosterone). If mutated get short plant. If replace with wildtype or human gene get plan of normal height. If overexpress the gene get large plant
Term
Cytokinins
Definition
cell division yielding shoot formation in culture, delay of leaf senescence, release of apical buds
Term
Ethylene (gas)–
Definition
Induced by auxin causes fruit ripening, root hair growth, abscission, senescence,
Term
Gibberillins
Definition
cell elongation, floral induction, seed germination.
Term
**Numb
Sensory Organ Precursor Cell (SOP)
neuron
sheath cell
Hair Cell
Socket cell
Definition
differentiated - glial mother cell (gets all of Numb)
neuroblast cell gets none of Numb

Body of adult fruit fly is covered with sensory bristles, each composed of a peripheral neuron, a hair cell, a sheath cell and a socket cell. These were derived from a sensory organ precursor (SOP) cell located in the surface epithelium of the pupa.

During pupal development the SOP undergoes 2 asymmetrical cell divisions. Numb goes to one of two daughter cells

IIb gets numb – divides into a sheath cell (no numb) and neuron (numb)

IIa does not get numb – it accumulates numb and divides into hair cell (gets numb) and socket cell (no numb).
Term
Inhibitory Signaling
Definition
A cell is destined to adopt a neural fate (has numb) it sends out inhibitory Information (delta and serrate). Through paracrine signaling to inhibit other cells. These cells have a notch receptor and are therefore prevented from being neural. Presence of Numb prevents the “neural fated” cell from receiving information in its notch receptor.

Homozygous notch mutants  lethal  all epidermal cells become neuroblasts
Term
Overview Segmentation
Hunchback, Nanos (inhibit hunchback - posterior), bicoid (anterior), Oskar
Definition
GAP genes – (mutations – large gaps in segments in Flies) are expressed just prior to cellular blastoderm stage. All encode transcription factors  MRNA’s proteins unstable short lived. Large areas anterior to posterior are specified. Then pair rule genes are expressed (early zygotic genes) specify 7 broad stripes.

Next get segment polarity genes (expressed during gastrulation) to subdivide 7 regions into 2 each. Therefore 14 narrower stripes, each with clear anterior and posterior polarity. Some transcription factors, some ligands, some receptors.

Finally homeotic genes, (homeotic selector genes) – type of homeobox genes specify the identity of the different segments
Term
Gap genes - Encode Factors Transcription
7 broad areas of the embryo
Turned on by zygote
Definition
Outlines the distribution of the Zygotically active gap genes; tailless, huckebein, giant, hunchback, kruppel, knirps. Gap genes establish 7 broad stripes in embryo; get very specific regions of gene expression. The genes may influence each other in complicated ways, i.e., bicoid (anterior) + hunchback anterior. Hunchback inhibits giant and therefore giant is found more posterior (no hunchback). However, bicord (anterior) is positive regulator of giant therefore even though hunchback is present still get giant
Term
Pair Rule Genes - Transcription factors
Definition
There are 8 pair rule genes – encode transcription factors

There are 3 - primary pair rule -> runt, hairy, eve (even skipped)

There are 5 – secondary pair rule -> ftz, opa, odd, slp, prd
Term
Regulation of Eve Expression
Definition
Early promoter for eve is influenced by gap genes and turns eve on in strips 2,3, & 7 (represented by sites E3, E7 and E2)

Maternal genes bicoid and hunchback activate and gap genes.
kr and gt repress

Eve expression -> Relationship determined if early enhancer eve is active. Hairy, runt, and eve needed to activate late enhancer to get eve expressed in other stripes and maintained in all.
Term
Segment Polarity Genes
Transcription Factors – Ligands Receptors
Definition
Segment polarity genes divide 7 regions from pair rule into 14 parasegments

4 key segment polarity genes engrail (en), naked (nkd) patched (ptc) and wingless (wg) (won’t discuss others)

Each is expressed in one cell of the 4 cell wide parasegment.

Pair rule gene eve positively influences engrail

Pair rule gene Ftz positively influences engrail
Term
Regulation of Engrailed
Definition
Expression of engrail (anterior of parasegment) stimulates hedgehog (Hh) production.
Engrail stimulate more engrail

Active Hh interacts with patched and smoothened (PTC-Smo) receptors on the adjacent cell (posterior end of earlier para segment) PTC inhibited wingless except when Smo is also stimulated therefore wingless (wg) is expressed.

Wg binds frizzled (a receptor on en expressing cell) stimulating En expression.

infulences itself and HH, which binds to both PTC and SMO, turns on WG. WG influences
Term
BX-C and ANT-C Complexes
Definition
Bithorax complex (Bx-C) -> 3 genes ultrabithorax (ubx), abdominal A and abdominal B (abdA & abdB). Null mutation Bx-C lose all abdominal segments and 3rd thoracic segment replaced by duplicated 2nd thoracic – lethal. UBx null mutation – T3 and A1 become T2 (flies with 4 wings) Ubx and AbdA, AbdB contain homeodomains which binds DNA (transcription factors). Mutations in homeobox -> homeobox mutations) (Not all homeoboxs are homeotic). Each gene has alternative promoters and splice sites -> very complicated regulation. Why? -> these genes repress anterior genes such as ant C complex genes

Gene complexes - must be expressed linearly in the embryo
Term
BX-C and ANT-C Complexes, continued
Definition
Arrangement Bx-Complex. If put Bx-C to another location get normal development. Rearrange order  major mutations. For some reason order is important in regulating those genes or their function.

Ant-C’s genes are arranged linearly, labial (lab), deformed (dfd-maxillary), sex comb reduced (scr) T1, and antennapedia (antp) T2, also gene pb (proboscipedia) all structures have T1 morphology – acts only in adults.

Null mutation – Labial parts of mouth become legs.
Term
7 segments into 14 parasegments of 4 cells
Definition
Gap rule genes influence Primary pair rule genes which influence segment polarity genes which influence homeotic selector genes
Term
BX-C and ANT-C Complexes, more
Definition
Gap genes -> stimulate pair rule gene expression -> stimulate homeotic gene expression
4 Homeotic genes -> Antp (from Ant-C) and the 3 BX-C genes are shown.

Posterior gene products prevent transcription of the more anterior genes (Blue Lines) and the proteins of these genes (red lines) allowing appropriate development.
Term
Anterior-posterior Patterning of Drosophila Wing
Definition
Wing development. Posterior wing cells express engrail proteins which stimulates (regulates) Hh and Hh is secreted. Anterior cells with patch and (partnered with) smoothened receptors bind Hh.

Stimulates Dpp (decapentaplegic) expression just anterior. Dpp diffuses anteriorly and posteriorly and organizes posterior and anterior portions of the wing.
Term
Dorsal-Ventral Patterning of Drosophila Wing
Definition
D/V interactions in wing development.

Apterous (Apt) and fringe (Fng) is 1) expressed dorsally and 2) Apt stimulate signal cascade between D & V compartments by stimulating dorsal cells to express serrate (membrane bound) ligand for the notch receptor.

For secreted protein fringe. Fringe and notch form a complex making notch less sensitive to serrate and increasing sensitivity to Delta (another ligand of notch) Localized expression of Fng binding to notch allows wingless to be expressed at the D/V border. Therefore, dorsally fringe secreting cells upregulate production of serrate due to Apt and lower sensitivity to serrate.

Ventrally -> sensitive to serrate. Activating notch at D.V. border stimulating wingless transcription which is needed for wing development. Wg gene produces WNT protein (wingless).
Term
Dorsal_Ventral Patterning of Drosophila Wing
Definition
The wingless protein (wnt) is activated by porcupine (porc) and binds to receptor family (frizzled)

Frizzled -> inhibits the destruction of Beta catenin.
Beta-catenin -> stimulates WNT response gene -> wing structure.
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