PROBLEMS in FRUIT BREEDING - POLYPLOIDY and HETEROZYGOSITY Polyploidy an Organism Having More Than Two Sets of Homologous Chromosomes Is Known As a Polyploid
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PROBLEMS IN FRUIT BREEDING - POLYPLOIDY AND HETEROZYGOSITY Polyploidy An organism having more than two sets of homologous chromosomes is known as a polyploid. Polyploidy is of general occurrence in plants while it is rare amongst animals. If the somatic chromosome sets in a diploid be represented by AA BB CC then the genome, i.e., the number in the genomes will be A B C. If this is represented by ‘n’ then the simple polyploid series would be: 2n – diploid 3n – triploid 4n – tetraploid 5n – pentaploid 6n – hexaploid 7n – heptaploid 8n – octaploid 9n – Nonaploid 10n – decaploid and so on Polyploidy is pervasive in plants and some estimates suggest that 30-80% of living plant species are polyploids, and many lineage show evidence of ancient polyploidy (paleopolyploidy) in their genomes. Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2n) gametes. Both autopolyploids (e.g. Potato) and allopolyploids (e.g. canola, wheat and cotton) can be found among both wild and domesticated plant species. Most polyploids display heterosis relative to their parental species. The mechanisms leading to novel variation in newly formed allopolyploids may include gene dosage effects (resulting from more numerous copies of genome content), the reunion of divergent gene regulatory hierarchies, chromosomal rearrangements, and epigenetic remodeling, all of which affect gene content and or expression levels. Many of these rapid changes contribute to reproductive isolation and speciation. Behaviour of polyploid crops Polyploid plants tend to be larger and better at thriving in early succession habitats such as farm fields. In the breeding of crops, the tallest and best thriving plants are selected for. Thus, many crops (and agricultural weeds) may have unintentionally been bred to a higher level of ploidy. The induction of polyploidy is a common technique to overcome sterility of a hybrid species in plant breeding. In some situations, polyploidy crops are preferred because they are sterile. For example, many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques such as grafting. Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine. Examples of polyploid crops • Triploid crops : banana, apple, ginger, watermelon, citrus • Tetraploid crops : potato, cabbage, leek, tobacco, peanut, kinnow, pelargonium • Hexaploid crops : chrysanthemum, bread wheat, triticale, oat, kiwifruit • Octaploid crops : strawberry, dahlia, pansies, sugar cane Some crops are found in a variety of ploidy. Apples, tulips and lilies are commonly found as both diploid and triploid. Bananas are available as diploid, triploid, tetraploid, and pentaploid. Daylilies (Hemerocallis spp) cultivars are available as either diploid or tetraploid. Kinnows can be tetraploid, diploid, or triploid. A survey of the chromosome numbers of the species in a genus or a family shows that these species generally fall into a polyploid series. The species are grouped together under a taxonomic head because of certain morphological resemblances and relationships. They may be crossable or may not hybridize at all with one another. However, the chromosome numbers of the species show a general relationship, i.e., they form multiples of a common basic number. The chromosome numbers of the family Solanaceae may be considered as an example. Types Autopolyploidy Autopolyploids are polyploids with multiple chromosome sets derived from a single species. They can result from a spontaneous, naturally occurring genome doubling, like the potato. Others might form following fusion of 2n gametes (unreduced gametes). Bananas and apples can be found as autopolyploids. Autopolyploid plants typically display polysomic inheritance, and are therefore often infertile and propagated clonally. Allopolyploidy Allopolyploids are polyploids with chromosomes derived from different species. Precisely, it is the result of doubling of chromosome number in an F1 hybrid. Triticale is an example of an allopolyploid, having six chromosome sets, allohexaploid, four from wheat (Triticum turgidum) and two from rye (Secale cereale). Amphidiploid is another word for an allopolyploid. Mango and banana are also allopolyploids. Doubled diploids are known as amphidiploids. Some of the best examples of allopolyploids come from the Brassicas, the three diploid Brassicas (B. oleracea, B. rapa, and B. nigra) and three allotetraploids (B. napus and B. juncea). Problems due to polyploidy and heterozygosity nature of fruit crops Fruit crops such as mango, banana and citrus pose the problem of polyploidy, and crops such as mango, papaya and citrus are highly heterozygous. Choosing of polyploidy varieties with desirable qualities may have the hindrance in developing hybrids as sometimes they exhibit sterility and obtaining a good hybrid may be questionable. In banana, when tetraploid is crossed with a diploid or triploid the genome of the segregating population will be unpredictable because of the restitution or unreduced chromosomes arising from the female parent. Heterozgosity on the other hand, create more complexity in breeding of mango, papaya and citrus because of wide segregations in the progenies. Hence, the breeding cycle is extended when compared to self pollinated crops because in every generation careful selection of progenies is required and high level of purity has to be maintained in each generation. Problems in Fruit Breeding – Polyembryony, Parthenocarpy and Seedlessness Polyembryony The phenomenon in which more embryos are present within a single seed is called polyembryony. It may result due to (a) nucellar embryony e.g., Citrus (b) development of more than one nucleus within the embryo sac (in addition to the egg embryo during the early stages of development) leading to multiple embryos (e.g. conifers). Occurrence of polyembryony is widespread in all citrus species but the number of embryos per seed varies from species to species. In rough lemon, it varies from 3 to 5. In mango certain cultivars are reported to be polyembryonic with the number of embryos ranging from 2 to 10 and the germination per cent from 40 to 87. Polyembryonic seedlings can be identified from its true seedlings by their uniformity and vigorous growth, while the seedling arising from fertilized embryo will be weak. The greater vigor in polyembryonic nucellar seedlings is probably due to the elimination of viruses. In mango polyembryony was determined by single dominant gene (Anon, 1996). In citrus, all the species are polyembryonic in nature except C.medica (Citron) and C.grandis (Pumelo) which are monoembryonic. Though nucellar embryony in citrus is of great value for producing vigorous, uniform and virus free plants, the phenomenon is an obstacle in hybridization. In polyembryonic cultivars, the vigorous growth of nucellar embryos inhibits the growth of the zygotic embryo and causes its degeneration prior to seed maturation. Such abortive embryos can be rescued by tissue culture. Parthenocarpy and Seedlessness In the recent years, the consumer preference towards seedless fruits is increasing among the consumers. The seedless nature of certain fruits is due to the phenomenon of ‘parthenocapy’ which refers to the development of fruits without fertilization or even without the stimulus that comes from pollination. Parthenocarpic fruits are usually seedless but need not be always. Vegetative parthenocarpy If a fruit develops even without the stimulus of pollination, then the phenomenon is referred to as vegetative parthenocarpy (automatic) eg. Banana and Japanese persimmon. Stimulative parthenocarpy If a fruit develops from the mere stimulus of the pollination (but without fertilization), the phenomenon is known as stimulative parthenocarpy. The female flowers of triploid watermelon require the pollen grains of diploid varieties to develop into a seedless fruit. Diploid pollen grain gives a stimulus to the ovary of guava when self pollinated, which result in the development of parthenocarpic fruit due to the stimulation provided by pollen hormones. E.g) Thompson Seedless variety of Grapes and papaya Steno-spermocarpy In “Black Corinth” variety of grapes, pollination and fertilization take place but the embryo gets aborted subsequently resulting in seedlessness. This phenomenon of development of seedless fruits is referred to as ‘steno-spermocarpy’. The seedlessness or parthenocarpic fruits are advantageous since there is a greater preference among the consumers for the seedless fruits of the same kind (e.g. seedless grapes, guava or oranges). Besides the problem of unfruitfulness due to pollination failure, sterility and incompatibility may not arise if a fruit develops parthenocarpically and the grower is assured of good crop (e.g. banana). One drawback with the seedless fruits is that they are usually small in size (e.g. Black Corinth variety of grapes) and irregular in shape (guava). Induction of seedlessness in fruits The seedlessness can be induced by the following methods. 1. Use of growth regulators Application of GA at 8000 ppm in lanolin paste on the cut end of the style of the emasculated flowers of guava resulted in the development of seedless fruits. Similarly, seedlessness in loquat was induced by spraying GA 100 to 200 ppm on the emasculated flowers. 2. Changing the ploidy level It was first demonstrated in Japan that by developing a triploid water melon 2n= 33 by