ANTERIOR POSTERIOR AXIS PATTERNING IN DROSOPHILA DROSOPHILA ORGANISATION. SEGMENTS C1, C2, C3 OF EMBRYO FORMS THE HEAD.

T1,T2,T3 FORMS THREE THORACIC SEGMENTS.T1 CARRY FIRST PAIR OF WINGS,T2 CARRY SECOND PAIR OF WINGS AND A PAIR OF WINGS,T3 CARRY PAIR OF HALTERES.THIS IS FOLLOWED BY 8 ABDOMINAL SEGMENTS(A1-A8)

Anterio-posterioir axis is laid down during oogenesis of Drosophila melanogaster.1 Maternal Effect Genes (MEG) play a malor role in patternng anterioposterior axis. MATERNAL EFFECT GENES Anterior Pole: Bicoid , Hunchback. Posterior pole: Nanos, Caudal. Bicoid mRNAs are attached to the microtubules and localized at the anterior end of oocyte. Nanos mRNAs are attached to the microtubules and localized at the posterior end of oocyte. The concentrations of Hunchback and Caudal mRNAs remain uniform along the anterior-posterior axis. Bicoid mRNA translated at anterior pole. Nanos mRNA translated at posterior pole. Bicoid protein inhibits translation of caudal mRNAs at anterior pole.Therefore caudal protein expressed only at posterior pole. Bicoid protein activates hunchback gene. Nanos protein blocks translation of hunchback mRNA at posterior pole.Hunchback protein expressed only at anterior pole. GAP GENES General regions of head, thorax and abdomen are mapped by the activity of these genes Involved in development of broad areas in embryos. First described by Christiane Nusslein Volhard and Eric Wieschaus in 1980s. Regulated by maternal effect genes. Important gap genes are kruppel, Knirps,hunchback,giant,huckbein,tailless.
1

Drosophila development is holometabolic ( Complete metamorphosis including four stages embryo, larva, pupa and adult(imago) stages).Beetles, moths, flies and butterflies exhibit holometabolic metamorphosis.

Kruppel is expressed mainly in the thoracic regions.Kruppel mutants lack thorax.

Gap genes code for transcription factors which regulate expression of pair rule genes. Gap genes . Pair rule genes, gap genes and segment polarity genes are segmentation genes. PAIR RULE GENES Involved in formation of individual segments.

Pair rule genes form individual segments

Important pair rule genes are fushi-tarazu, even-skipped, hairy, oddskipped, paired, runt Fushi-tarazu is expressed in the boundaries of segments.

fushi-tarazu is expressed in borders of segments.

After gastrulation 14 parasegments are produced actually.Posterior half of one parasegment and anterior half of adjacent parasegment forms a segment. Mutations in pair rule genes causes loss of alternate segments.Phenotype:number of segments reduced to half.

SEGMENT POLARITY GENES Gives each segment an anterior/posterior orientation.

segment polarity genes in action

Mutation in segment polarity genes:n change in number of segments.Parts of segments affected. Engrailed is a segment polarity gene. In engrailed mutants the posterior end is replaced by a mirror image of anterior end.

Acts in parasegments(14 in number) By the time segment polarity genes are active the embryo would be cellularised(cellular blastoderm).Therefore segment polarity genes act in cells.(NOT SYNCYTIUM) HOMEOTIC GENES Specifies the types of appendages formed on each segment. They code for transcription factors. The genes carry a 180bp long sequence called the HOMEOBOX. The homeobox encodes for a 60 amino acid region called the homeodomain. Homeodomain confers DNA binding function. Homologues of homeotic sequences found in insects,nematodes,mollusks,fishes,frogs,birds etc. Two classes of homeotic genes Bithorax complex and Antennapedia complex. BITHORAX COMPLEX(3-GENES). -Ultrabithorax(ubx), abdominal(abd-A),abd-B -ubx is expressed in T3 and A1. -expression of ubx in T3 prevents it from becoming a T2 -ubx mutation results in conversion of T3 to T2 resulting in formation of 4 wings.Such flies cannot fly. ANTENNAPEDIA COMPLEX. -Antennapedia normally expressed in T2 -if expressed in head causes head to develop legs.