KYRANCONNCE

NCEPT OF GENE (Structural Organization of Gene)

MOOERN
311
c-onc gene, but the v-onc genes
the
uninterrupted coding sequences, while Homeoti genes
situated on a
are
enes have thè usual genomic orga- Segnent of
chromoome, may be as particular
exon and intron.
as cluster of
cated genes which interact mutigene
in
or
t of alternating compli-
interloxking pattern that is not fully defined a

and
cellular oncogenes otooncogenes Many homeotic
hane'.
multiple exons
separated by introns, where- factors that act upongenes code for
other hormetic transcription
s t h e viral
oncogenes are single exons. For as
upon other target loci. As a result, genes as well
chicken cellula src protooncogene one horneotic gene may a mutation in
e.mple,

11 introns separating 12 coding influernce

of other hormeotic genes. The the expression
contains

whereas e RSV
the RSV V-sr
v-src gene has a sin- tion of sorne hormeotic molecular organiza
guences, rise to their genes is unusual
and gave
uninterrupted coding sequence. Both the description as complex loci, and itself
de for protein
kinases that phosph0 poses some
questions about gene expression. The
enes

tyrosine residues, oth the protein can homeotic mutants develop organs at wrong
yiate same antibody. It has been found due to places
inability to recognise their positional intfor
eract with
Dlerac

thatthere
are 18 single
eotide pair diffe-
es between the coding sequences of v-src
nd C-SIC that result in 8 amino acid changes
but those must not
te protein products,
involved in oncogenecity.
in several cases, the v-onc coding sequence
appears to have evolved only relatively little from
the c-onc sequence, usually by point mutations.
An indication that the v-onc genes retain func-
tions related to their c-onc progenitors, is provi-
ded by the fact that the majority of nucleotide
changes are located in third base positions where
they do not afect the protein sequence.
So, in a word- viral oncogenes and cellular
oncogenes are almost same, but the cellular
Oncogenes are present as protooncogene. These
prOtooncogenes can mutate to form that are
capable of inducing oncogenesis.
HOMEOTIC GENES
mation properly, as a result the organs thermselves
were normal but their
developed were improper. positions_at which they
in
be Embryogenesis in Drosophila
Homeotic
genes are involved during embryogenesis in
Drosophila. Their expression depends on the
prior expression of segmentation genes. The seg-
mentation genes define the number and loca-
tions of segments, and homeotic genes impose
the programme that determines the
unique
differentiation of each segment. So homeotic
mutants cause the transformation of a segment to
develop phenotype of other segment, i.e. legs
may develop in place of antennae or wings may
develop in place of eyes, etc.
The classic complex homeotic locus is
BX-C, the bithorax complex, characterized by
several groups of homeotic mutations that affect
development of the thorax, causing major
morphological changes in the abdomen (Fig.
14.26). Another such example is ANT-C(anten
napedia complex), aftecting head and thorax.

Homeotic genes constitute an intricate regu-

latory
These homeotic genes, which
the
were also

network, in which one gen may activate called Hox genes, are characterized by
repress other enes. Homeotic genes are presence of a homeobox
of 180 nucleotide
Involved in regulation of develop through a sequence,
translated into 60 amino acids
actual-
termined domain called homeodomain. helix-turn-helix
These are

turned pathway
are in which specific genes
with
on and off at particular times. ly DNA binding protein
Homeot
in whieic genes
es act in a transcriptional cascade
ch a series
(HTH) contormation playing regulatory
role for
at major
of hierarchical interactions transcription of DNA by binding
DNA double helix. However, Hox
the n the regulatory proteins is succeded by groove of
subset of homeobox
tivation of ctural genes coding for genes represent
only a

morphogenesis. genes, since many

box may not be
genes containing
homeotic in nature.
a homeo-
 CFLL BIOLOGY, GENETICS AND MOLECtR
12 JLAR BIOCK
CARPELS (AGC
ST
AMENS AG d

PETALS
SEPALSAPA
APA
Sepals Petals Stamens Carpels
Fig. 14.26: Afour-winged fly is produced by a triple muta
tion in abx, bx, and pbx at the BX-C complex. AP3
For details of BX-C locus, see Fig. 14.13]
AP1
AP2 AG
SEP1
Flower Primordia Development:Induction SEP2
of tower development in plants takes place by SEP3
the changes from vegetative to reproductive Category A :AP1 andAP2-
Sepal and Potal
meristems or floral primordia and homeotic Category B:AP3 and P- Petal and gimora
genes act sequentially controlling development. Category C :AG-Stamen and Carpel
Stamen primori
Flower development requires a cascadeof primordla
Fig. 14.27: Floral homeotic
sequential gene activity that gradually converts a Simpson) genes-categories
mass of undifferentiated apical meristem cells
into the different parts of a flower. The
genes
encode transcription factors that act as master carpel, it is repressed by the repressor
whic
another gene product
switches turning on or off downstream genes that of CURLY
LEAF
ultimately make each part of the flower in its In floral development, the class A
class gene a
appropriate location. gene are believed to
function antagons
The mechanism of flower morphogenesis in tically, because, in a mutant (ap2) of class
Arabidopsis has been deduced through a genetic gene, the expression
clas C (AG) ge of
model called ABC model (Coen and expanded to whorls one and two. In mutant a
1991). The
Meyerowitz, of class C gene, AP1 expands its
floral homeotic genes have been expression
studied and classified into 3 the inner whorls. The expression of Pl
and
categories -

A, B and AP
C. Genes from I.e., class B genes is independent of class A
category A
APETALA1 (AP1)
-

and APETALA2 (AP2) are expressed in sepal and C, but Pl activity is required for AP3 expres"
petal primordia; genes from and vice versa (Table 14.2).
category B -

APETALA3 (AP3) and

PISTILLATA (PI) are Homeotic genes have also been identiet
expressed in petal and stamen
primordia; and other plant systems Iike Antirrhinum (LIP D
gene from category C AGAMOUS (AG) affect GLO, PLE, FAR), Petunia (GP, BL, DOL, F
stamens and carpels (Fig.
the revised ABCE 14.27))According
model, another category (E) of
to Gerbera (GRCD), etc.
genes SEPALLATA (These iloral organ identity genes have
with A, B, C (SEP1-4) act in combination shown to encode transcriptionfactor each
genes for specification of floral MACN
organs. The LEAFY (LFY) gene identified, is conserved MADS-DNA binding domain inital
responsible for specifying floral meristem identi-
domain has 56 amino acids which was factr

ty and plays a major role in found in 4 different proteins: transcrip

groups of genes for floral turning
on the above
MCM of yeast, proteins encoded by AGAs
After expression of the organ development. and DEFICIENS genes of Arabidop
gene AGAMOUS (AG), man
involved in the Antirrhinum, ription factor ofSRf
O
development of stamen and mais; and from these four proteins ue
nare
 OF GENE
(Structural Organization of Gene) 313
O N C E P T

p E R NCON

organ
identity genes in abidopsis thaliana (from P. K. Gupta)
k2
Floral
Gene and its symbol Mutant allele and expression
s sOl
g e n e

APETALA1 (AP1) ap1 (Sepals replaced by bract like structures

A and petals aborted or sepaloid)
APETALA2 (AP2) ap2 (Sepals replaced by carpelloid structures
and petals replaced by stamens)
APETALA3 (AP3) ap3 (Petals replaced by sepals and stamens
B replaced by carpels)
PISTILLATA (PI) pi (Similar to above)
AGAMOUS (AG) ag (Stamens replaced by petals and carpels
C replaced by sepals)

has been originatedThis MADS domain from mutant analysis of amino acid biosynthetic
MADS pathway of Neurospora, and biochemical
muta-
binding domain and also helps
in
DNA tions concerned with the metabolic disease of
interactions. The homeotic MADs
agtein-protein human. The concept of one gene-one polypep
a subset of a larger MADDS
box genes are actually of whose members are
tide emerged from the study of haemoglobin
box gene family,
some
from
whereas others show polypeptides in human beings suffering
expressed in all tissues, sickle cell anaemia.
or organ specific expression.
fiSsue
Position effect as shown in Bar locus of
Many MADS box genes have been cloned
various plants and expression patterns
Drosophila indicated that gene is not a point, but
from in has its dimensions. Closely linked genes, having
show the common gene regulatory system
similar phenotypic effect, behaving ordinarily as
flower development. De Folter et al. (2005) have
re-occurring alleles, but be separable by crossing over are
Concluded that self regulation is a
termed Pseudoalleles, e.g., lozenge locus in
phenomenon among the MADS domain pro
ens. Since many single MADS domain proteins Drosophila.
can form complexes with several other
MADS pseudoallelism or cis-trans effect
Position
domain proteins, the number of possible explains the lack of complementation in F indi-
Combinations is very large and may permit
the viduals. cis-trans complementation test is used to
contexts. The determine the functional allelism of any two
proteins to function in different AG recessive genes.
protein,
Tunctional view of, for example, the identity, 1s The concept of ultimate unit of heredity for
determining stamen and carpel
AG interacts with different sets
of MADS the genes, underwent modification following the
ut func- work of Benzer on bacteriophage. On the basis
0main factors to carry out the different
tions. of complementation test and recombination test,
he coined the terms cistron the unit of function
(subdivision of gene), recon the unit of recom-
SUMMARY certain
bination (subdivision of cistron), muton - the unit
of mutation (subdivision of recon). The
materials, have genes of
the hereditary the classical authors are comparable in structure
Enes, autocatalysis, heterocata-
ntial properties to that of cistron.
classical concept gene of
and mutation. The
Is molecular
revolutionized by Genetic fine structure maps have been
been con-
dSgreatly structed for many genes by deletion
Concept in recent years. is mediated
obtaining a series of partially mapping. By
deletions, any point mutation can beoverlapping
characters
control of all relationship
uenetic The
gh specific enzymes
is 1:1
as
evidenced
testing the ability to recombine mapped by
etwee cEen gene and
enzyme
among them.