Cross-breeding
Cross-breeding is another well-known means of genetic improvement which
has application in aquaculture. Heterosis or hybrid vigour enables an offspring
to surpass its parents for one or more traits. On the other hand, inbreeding
depression caused by mating of closely related parents has a deleterious effect
on the progeny.
The inbreeding measure is the coefficient of inbreeding, incorporating
the degree of the animal’s homozygosity. It shows what part of the genes in a
group of individuals are in the homozygous state. Close inbreeding, especially
sib mating (brothers and sisters and parents with offspring), causes
homozygosity to increase rapidly, to as much as 0.9–0.95 (expressed as
fractions of one) or even more. In most animals, inbreeding results in
inbreeding depression, characterized by a drop in viability, growth rate and
often fertility. Such depression has been observed by many workers in common
carp, brook trout and other fish species. Outbreeding, on the other hand, is
accompanied by heterosis in growth rate and viability, especially when fish
from different high inbred groups are crossed. The main types of crossings
undertaken are:
a)
Commercial crossing, directed
towards breeding of the first-generation hybrids for commercial purposes. Only
the first generation, that has the heterosis of productive qualities or
incorporates the advantageous characteristics of both the parental forms, is
used. They are maintained for further reproduction.
b)
Synthetic or distant crossing, in
which distant parents, including those of inter-generic origin, are crossed to
develop a new breed, in the course of long selection. It may attempt to combine
the qualities of parents of several breeds, species or even genera. Such
crossings to produce new breeds should ensure the preservation and perfection
of the productive qualities of the breed, the preservation of genetic
variability and the prevention of inbred depression.
Distant outbreeding is indispensable in the selection of aquaculture species.
The aims of such crossings are as follows (Kirpichnikov,
1971):
(1) an overall increase in genetic variability, resulting in
an increase in
selection response;
(2) achievement of a combination of charac teristics of two or three
breeds or two (rarely three) species;
(3) improvement of the productive quality of the local breed by making
use of the few valuable traits of another breed;
(4) increase in the viability of the breed by intro ducing genes
responsible for resistance to environmental factors and diseases.
Kirpichnikov (1971) describes different cross- breeding patterns to
achieve these aims. Reproductive
crossing is suggested
when valuable properties from both parents are to be combined in the
hybrids. It can be done with
complete fertility of the hybrid and requires only meticulous selection
in the subsequent generations.
Introductory crossing will be advantageous when one or only a few
characteristics from a breed have to be incorporated in the hybrid. Each
generation of the hybrid has to be crossed with the local breed and so there is
the risk of losing the useful characteristics of the improved breed in back
crossing, particularly in the case
of polygenic inheritance of properties selected.
This type of crossing is of considerable use in selection for increased
resistance to certain dis- eases, which is often dependent on the presence of
one or a few genes. These genes can be pre served by means of proper selection
in each generation.
Absorptive crossing differs from introductory crossing only in that its
purpose is a nearly com plete substitution of the local breed of geno- type by
the genotype of the improved breed. Only some features of the local breed, such
as
viability, are preserved.
Alternate crossing is the most complicated system which is most useful
when a combina tion of many characteristics from two breeds with polygenic inheritance
is required. It allows the preservation of
high genetic variability through a number of generations.
Selection effi ciency is kept at a high level owing to this variability and
does not reach a plateau. The mainproblem of obtaining new hybrid breeds by means
of crossing (interspecific or intergeneric) is their complete or partial
sterility, which takes a lot of time-consuming work to overcome.
A number of breeding systems have been proposed to utilize completely
the advantages associated with heterogeneous crossings. Paral lel breeding of
two or more groups within a breed is possible when working with slowly maturing
species, without intermingling, allow ing a moderate inbreeding among each and carrying
out selection in each generation. In breeding in groups for family selection, a
large number of crossings from different groups are carried out for each
generation. The parents producing the best offspring are used for subsequent commercial
crossings. This systemsuffers from the drawback that the
genetic variability gradually decreases during family selec tion. Moav and
Wohlfarth (1967) recommend that a reserve group of a sufficient number of individuals
should be kept for each group when selecting two groups marked by certain
genes.
In case of a drop in genetic variability, addi tional gene pools can be
introduced into the exhausted group.
Another possible system
is alternate in- breeding and outbreeding. After two or
three generations of close inbreeding, the evaluation of hybrids from different
inbred lines is per formed. The best combinations are used for
commercial rearing and among the offspring new inbred lines are
established. Linear selec tion involving inbreeding for superior ancestors and
top cross, where crossing is done between the best inbred individuals (say
males) and individuals from the outbred population (say females) to preserve
the genetic variability, are
other methods that
have applications in aquaculture.
One of the
most complicated techniques of breeding is reciprocal
recurrent selection (RRS), where the combining capacity of the parents from
each of the two groups is evaluated by means of a cross with parents from the
other group. The individuals thus selected are reproduced without
recrossing and their off- spring again tested for combining potential. The
basic feature of all the systems described is
the utilization of
heterosis in crossing
individuals from different groups, lines and breeds. Along with this,
moderate to very close
inbreeding is employed. The most appropriate system would naturally
depend on the species and the traits that are of importance in commercial
culture. Gjedrem (1985), however, proposed a cross-breeding scheme for fish
farming as summarized below:
a)
Test all possible crosses between
different strains or species for the economic traits in question and select
crosses that are likely to give useful results. It may be better to use strains
with very different origins and which, in combination, have favourable traits.
b)
Develop inbred lines and test the
crosses under natural conditions to find the most valuable cross for farming.
c)
Start an RRS programme to ensure
continuous genetic improvement, utilizing both general and specific combining
abilities simultaneously.
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