Sexual incompatibility Stigma receives a variety of air-borne and insect carried pollens.

However, only a few succeeds in effecting fertilization. Pistil is equipped with devices to allow the pollen of only the right mating type to function normally, others are discarded. If a pistil carrying functional female gamete fails to fertilize with viable and fertile pollen, capable of bringing about fertilization, the two are said to be incompatible. The phenomenon is known as sexual incompatibility. Sexual incompatibility may be divided into (a) interspecific (between individuals of different species), (b) intraspecific (between individuals of the same species) also called self-incompatibility. Interspecific incompatibility is heterogenic, i.e. controlled by more than one gene at different loci on the chromosomes. It prevents cross-pollination. It may be caused by (a) Either fertilization does not occur, (b) syngamy is followed by abortion of hybrid embryo due to inadequate development of endosperm or (c) embryo-endosperm incompatibility. Self-Incompabiility Many flowering plats are outbreeders which meant that they are successfully fertilized only by the pollen of other plants and not by their own. In nature, different floral adaptations, such as dichogamy, herkogamy and unisexuality have evolved to prevent self pollination. However the most effective natural device to enforce outbreeding is self incompatibility. Self incompatibility can be morphologically divided into two types : (a) Heteromorphic : It is characerised by the occurrence of two (distyly) or three (tristyly) morphologically distinct mating types within a species which can be easily recognized without a breeding test. The difference generally occur in the position of stigma and anthers (heterostyly). In these systems, the incompatibility reaction may be controlled by one gene with two alleles, (Primula) or by two genes with two alleles each (Lythum). Other features are a. The incompatibility reaction is determined sproophytically b. The growth of incompatible pollen tubes is inhibited in the style c. Dominance between alleles of the incompatibility genes is expressed both in the pollen and the style. (b) Homomorphic : In this tpe, all the mating types within a species are morphologically indistinguishable and require proper breeding test for recognition. A species with this type of incompatibility has numerous mating types. On the basis of origin of factors determining mating types, two categories of self incompatibility have been recognized : (a) Gametophytic self incompatibility (GSI) : the incompatibility is determined by the genotype of male gametophyte itself e.g. Gramineae, Liiaceae, Solanaceae, Trifolium. It is common in species with 20celled pollen. Pollen usually germinate normally and inhibition occurs at the style. (b) Sporophytic self incompatibility (SSI) : the incompatibility is controlled by the genotype of the sporophyic tissue of the plant from which the pollen is derived, e.g. Compositae, Cruciferae. It is common in species with 3-celled pollen. Inhibition occurs at the stigma. Genetic Basis of Self Incompatibility Most popular hypothesis about the genetic control of self incompatibility is Opposition S-alleles first proposed by East and Mangelsdorf (1925). According to this, incompatibility reactions are controlled by a single gene, called Sgene, which has several alleles. In GSI, pollen grains that possess the S-allele common to any one of the two alleles present in the cells of the pistil will not be functional on that particular pistil. (S1S2)pollen (S1S2) pistil no fertilization. (S1S2) pollen (S2S3) pistil 50% fertilization. (S1S2) pollen (S3S4) pistil 100% fertilization. The S-alele of the pollen or the male gametophyte, whih determines the incompatibility reaction. The inhibition occur at the stylar canal of the pistil. In SSI, all the pollen of a plant behave similarly, irrespective of the S-allele they carry. A plant carrying S1S2 alleles the pollen carrying S1 or S2 would behave as S1 or S1 plus S2 (if no dominance. The presence of one of the alleles of the stylar tissue in the sporophytic tissue of the male parent would render all the pollen of that plant non functional with respect to that particular style.

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(S1S2) pollen (S1S2), (S1S3), (S1S4), (S2S3),(S2S4) and so on pistil no fertilization. (S1S2) pollen (S3S4), (S3S5) pistil 100% fertilization. Barriers to fertilization : In incompatibility systems barriers to fertilization may occur at different levels : (a) Pollination : methods adopted by plants are to check illegitimate self pollination, such as dichogamy, herkogamy and unisexuality. (b) Progamic phase : inhibition occurs at any stage from pollen germination to the discharge of male gametes in the vicinity of egg. a. On the stigma : pollen fails to germinate or the pollen tube cannot penetrate into the stigma (e.g. Cruciferae, Gramineae) b. On the style : pollen tubes fail to grow through the full length of the style (e.g. Liliaceae, Solanaceae). (c) Syngamy : pollen tubes fails to enter the ovary (Hemerocallis), or its growth is arrested within the ovule (Acacia) or within the embryo sac (Theobroma). Male and female gametes fails to unite. Physiology and Biochemistry of Incompatibility The incompatibility process are controlled by two reactions : (a) Recognition reaction : it occurs at the molecular level and by this the pistil recognizes whether a particular pollen grain is to be accepted (compatible) or rejected (incompatible) (b) Rejection reaction : It sets in physiological and biochemical process in the pistil specific to the type of pollen received. In incompatible pollination, the phenotypic manifestations of these changes bring about the rejection reaction. It may occur on the stigma or prevent the entry of pollen tube into the stigma (SSI) system or it may occur in the style inhibiting pollen tube growth (GSI system). There are some role of pollen wall and the pollen proteins in the pollen-stigma interactions. Pollen grains on hydration released some hydrolytic enzymes, which played significant role in the pollen-stigma interaction. The inter and intra-specific incompatibility is associated with recognition proteins held extracellularly in the pollen wall. These derived proteins are released on hydration of the pollen. Stigma surface inhibition : In this type, recognition and rejection occur on the stigma. If the pollen which fails to germinate on the stigma is introduced directly into the style, it causes normal fertilization. It shows that stigma is the barrier of pollen germination. In Cruciferae, the stigma is covered with a cuticle. If the pollen can penetrate the cuticle, it is compatible. If the pollen fails to grow through the cuticular layer, it is incompatible. Compatible stigma activates the enzyme cutinase. Recent studies show that (a) Rejection reaction exhibited by incompatible stigma lack copious exudates i.e. remain dry (b) Pellicle comprises of lipid containing a variety of proteins which possess esterase activity (c) Pollen stimulates exudation of water by the papillate cells. In turn, stigma stimulate the outflow of exine bound proteins. (d) The released exine proteins are received by the pellicle where the interaction occurs in the stigma. (e) In incompatible pollen, callose plugs appear just below the pollen contact point. Stylar inhibition : In GSI, the pollen germinates, and grows into the style. Then the rejection reaction sets in stopping the pollen tube growth. Stylar inhibition is based on the following characteristics (a) The cells of the style secrete S-allele specific proteins which induce incompatibility reactions (b) Probable involvement of peroxidase in rejection reaction (c) Degradation of tube cytoplasm and thickening of the walls of the pollen tube (d) Tip of the pollen tube with incompatible styles show different structure and unable to absorb the secretion of the pistil (e) Styles of self incompatible species exhibit high RNAse activity causing checking the growth of the pollen tube. (f) Abnormal behavior like branching of the tip of the pollen tube, development of callose plugs or restricted growth of pollen tube. Biological Significance : In nature a balanced inbreeding and outbreeding of the plants is regulated by intraspecific and interspecific incompatibility. Selfing of the plant lead to homozygous individuals which have a very low survival value. To prevent this, nature has imposed self incompatibility. In interspecific incompatibility rings about reproductive

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isolation and is responsible for the emergence of new races and species through the process of mutation and open hybridization. On the other hand, incompatibility may be a hurdle in plant improvement programme. Methods to remove sexual incompatibility 1. Pollination methods : modification in the pollination patterns a. Bud pollination b. Intra-ovarian pollination c. Mixed pollination d. Stub pollination e. Test tube pollination 2. Irradiation method : inducing cytological changes in the pollen or remove the inhibitory substances in the style 3. Modification of the stigma : by applying NaCl to the stigmatic suface or other compounds. 4. Heat treatment of style 5. Parasexual hybridization : three steps a. Isolation of protoplasts b. Fusion of isolated protoplasts c. Culture of protoplasts.

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