ARUN KUMAR
ASSISTANT PROFESSOR (HOD)
DEPARTMENT OF ZOOLOGY
A.N.S. COLLEGE, NABINAGAR
AURANGABAD
LINKAGE AND CROSSING
OVER
LINKAGE CAN BE DEFINED AS THE TENDENCY OF GENES
TO REMAIN TOGETHER IN THEIR ORIGINAL
COMBINATION DURING INHERITANCE IS CALLED
LINKAGE.
MENDEL’S LAW OF INDEPENDENT ASSORTMENT STATES
THAT THE SEGREGATION OF ALLELES OF ONE GENE IS
INDEPENDENT OF ALLELES OF ANOTHER GENE. THIS
LAW ALWAYS HOLDS TRUE FOR GENES THAT ARE
LOCATED ON DIFFERENT CHROMOSOMES, BUT FOR
GENES THAT ARE ON THE SAME CHROMOSOME, IT DOES
NOT ALWAYS HOLD TRUE.
GENES PRESENT ON A SINGLE CHROMOSOME IS SAID TO BE
BELONGING TO ONE LINKAGE GROUP. THEREFORE,THERE ARE AS
MUCH LINKAGE GROUP AS THE NUMBER OF HAPLOID
CHROMOSOMES.
LINKAGE WAS DISCOVERED THROUGH A CLASSIC EXPERIMENT BY
WILLIAM BATESON AND REGINALD PUNNET.THEY WERE STUDYING
THE INHERITANCE PATTERN OF THE GENE FOR FLOWER COLOUR
(P-PURPLE AND p-RED) AND THE GENE AFFECTING THE SHAPE OF
POLLEN GRAINS (L-LONG AND l-ROUND).
THEY CROSSED THE PURE LINES PPLL AND ppll AND THEN SELF
CROSSED THE RESULTING PpLl LINES. THE OBSERVED FREQUENCY
DIFFERED FROM THE EXPECTED 9:3:3:1 RATIO OF PL:Pl:pL:pl. THEY
OBSERVED AN INCREASED FREQUENCY OF PL AND pl AND A
DECREASED FREQUENCY OF Pl AND pL .
CROSS SHOWING LINKAGE
CROSS SHOWING LINKAGE
THEIR EXPERIMENT REVEALED LINKAGE BETWEEN THE P AND L
ALLELES AND THE p AND l ALLELES.THE FREQUENCY OF POCCURING
TPGETHER WITH L AND WITH p OCCURING TOGETHER WITH l IS
GREATER THAN OF THE RECOMBINANT Pl AND pL.
GENES PRESENT ON THE SAME CHROMOSOME IS THEORETICALLY
LINKED TO EACH OTHER, BUT MOST OF THE TIME THEY SHOW
INDEPENDENT ASSORTMENT AS OBSERVED BY MENDEL. THIS
PHENOMENON IS DIRECTLY RELATED TO THE DISTANCE
BETWEEN THE TWO GENES IN QUESTION. THE MORE THE
DISTANCE BETWEEN THE TWO GENES THE GREATER
PROBABILITY OF THEIR INDEPENDENT ASSORTMENT, THAT IS
THEY WILL BE SEPARATED DURING GAMETE FORMATION. THIS
PHENOMENON IS FACILITATED BY THE PROCESS CALLED
CROSSING OVER.
CROSSING-OVER CAN BE DEFINED AS THE EXCHANGE OF
GENETIC MATERIAL BETWEEN THE NON-SISTER
CHROMATIDS OF A HOMOLOGOUS CHROMOSOME. THE
MOST IMPORTANT FEATURES OF CROSSING OVER CAN BE
SUMMARIZED AS FOLLOWS:
o THE GENES ARE LOCATED ON A CHROMOSOME IN A
LINEAR SEQUENCE AT
DIFFERENT LOCI.
o THE TWO ALLELES OF A GENE IN A
HETEROZYGOTE OCCUPY CORRESPONDING POSITIONS
IN THE HOMOLOGOUS CHROMOSOMES.
o CROSSING-OVER INVOLVES THE BREAKAGE OF EACH
OF TWO NON-SISTER CHROMATIDS OF THE
HOMOLOGOUS CHROMOSOMES
CROSSING-OVER OCCURS AT PACHYTENE STAGE AFTER SYNAPSIS OF
THE HOMOLOGOUS CHROMOSOMES HAS OCCURED IN PROPHASE 1
OF MEIOSIS.
CHROMOSOMES WITH RECOMBINANT COMBINATIONS OF LINKED
GENES ARE FORMED BY THE OCCURRENCE OF CROSSING-OVER IN
THE REGION BETWEEN THE TWO LOCI.
A CROSS-SHAPED STRUCTURE IS FORMED DURING CROSSING-OVER
IN WHICH TWO OF THE FOUR CHROMATIDS OF HOMOLOGOUS
CHROMOSOME PAIRS APPEAR TO EXCHANGE MATERIAL. THIS
STRUCTURE IS CALLED CHIASMA.
TH MORGAN WAS THE FIRST TO HYPOTHESIZE THE RELATION
BETWEEN LINKAGE AND SEGREGATION OF HOMOLOGOUS
CHROMOSOMES AND THE OCCURRENCE OF CROSSIONG-OVER
BETWEEN HOMOLOGOUS CHROMOSOMES DURING MEIOSIS.
CROSSING OVER BETWEEN HOMOLOGOUS
CHROMOSOMES
TYPES OF LINKAGE
GENERALLY IT IS OF THREE TYPES---
01.CROSSING-OVER
02.GENES INVOLVED AND
03.CHROMOSOMES INVOLVED.
01.BASED OF CROSSING OVER
IT IS OF TWO TYPES:-
(A)COMPLETE LINKAGE:-IT IS KNOWN IN CASE OF MALES OF
DROSOPHILA AND FEMALES OF SILKWORMS,WHERE THERE IS
COMPLETE ABSENCE OF RECOMBINANT TYPES DUE TO ABSENCE OF
CROSSING OVER.
(B)INCOMPLETE /PARTIAL LINKAGE:-IF SOME FREQUENCY OF CROSSING
OVER
ALSO OCCURS BETWEEN THE LINKED GENES, IT IS KNOWN AS INCOMPLETE
/PARTIAL LINKAGE. .IN COMPLETE LINKAGE HAS BEEN OBSERVED IN MAIZE,
PEA, DROSOPHILA FEMALE AND SEVERAL OTHER ORGANISMS.
02.BASED ON GENES INVOLVED
BASED ON GENES INVOLVED :-DEPENDING ON WHETHER ALL
DOMINANT OR SOME DOMINANT AND SOME RECESSIVE ALLELES ARE
LINKED TOGETHER ,LINKAGE CAN BE CATEGORIZED INTO
(a)COUPLING PHASE AND
(b)REPULSION PHASE.
(a)COUPLING PHASE:-DOMINANT ALLELES PRESENT ON THE SAME
CHROMOSOME AND RECESSIVE ALLELES PRESENT ON SAME
CHROMOSOME SHOWS COUPLING PHASE
TR tr
----- ----- COUPLING PHASE
TR tr
(b)REPULSION PHASE:- DOMINANT ALLELES OF SOME
GENES ARE LINKED WITH RECESSIVE ALLELES OF OTHER
GENES ON SAME CHROMOSOMES SHOWS REPULSION
PHASE
Tr tR
--- ---- REPULSION PHASE
Tr tR
03.BASED ON CHROMOSOMES INVOLVED:-
BASED ON THE LOCATION OF GENES ON THE CHROMOSOMES,
LINKAGE IS CATEGORIZED INTO
(a)AUTOSOMAL LINKAGE:-IT REFERS TO LINKAGE OF THOSE GENES
WHICH ARE LOCATED IN AUTOSOME (OTHER THAN SEX
CHROMOSOMES)
(b)ALLOSOMAL LINKAGE /SEX LINKAGE:-IT REFERS TO LINKAGE OF
GENES WHICH ARE LOCATED IN SEX CHROMOSOMES i.e. EITHER “X”
OR “Y”(GENERALLY “X”).
LINKAGE GROUPS
LINKAGE GROUP REFERS TO A GROUP OF GENES WHICH ARE
PRESENT IN ONE CHROMOSOME. IN OTHER WORDS ALL THOSE
GENES WHICH ARE LOCATED IN ONE.
CHROMOSOME CONSTITUTE ONE LINKAGE GROUP. THE NUMBER OF
LINKAGE GROUPS IS LIMITED IN EACH INDIVIDUAL. THE MAXIMUM
NUMBER OF LINKAGE GROUPS IS EQUAL TO THE HAPLOID
CHROMOSOME NUMBER OF AN ORGANISM.
E.g.- FOR EXAMPLE THERE TEN LINKAGE GROUPS IN CORN (2n=14),
FOUR IN DROSOPHILA MELANOGASTER (2n=8) IN MAN (2n=46).
THANK YOU
