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Review of Literature REVIEW OF LITERATURE The present review deals mainly with studies on various aspects of root nodule bacteria associated with wild legumes. However, whenevernecessary, relevant literature regarding studies on similar aspects of other cultivated legumes and their bacteria has also been incorporated. The review has been divided into the following parts 2.1 NODULATION STATUS 2.1.1 Black-coloured nodules. 2.2 STUDIES ON ROOT NODULE BACTERIA 2.2.1 Classification 2.2.2 Cultural and Morphological characteristics 2.2.3 Biochemical and Physiological characteristics. 2.3 PLANT INFECTIVITY AND CROSS-INOCULATION STUDIES 2.3.1 Plant infection test 2.3.2 Cross-inoculation studies 2.3.3 Nodulation test on Siratro 2A CROP RESPONSE TO INOCULATION 2.4.1 Competition studies 2.if.2 Application of rhizobia isolated from wild legumes on cultivated legumes. 2A.3 Evaluation of symbiotic nitrogen fixation efficiency. 2.5 NITROGEN FIXATION BY WILD LEGUMES. 2.1 NODULATION STATUS The ecological uniqueness of the Leguminosae derives from the tubercles or nodules of their root systenns. A wide variation is seen in the nodulation ability of leguminous plants. Allen and Allen (1981) who have compiled extensive nodulation survey data of Leguminosae, claim that ^8% of leguminous genera have been examined for nodulation status. Out of this percentage, 84% were found to be nodulated. Of the genera examined in Mimosoideae and Papilionoideae, 83.87 % and 94.17 % respectively included nodulated species, whereas in Caesalpinoideae only 40% of the genera examined showed nodulation ability. Although tropical flora is rich in leguminous plants, there is very limited knowledge about the nodulation of tropical legumes (Banadoz and Fernandez, 1954; Bowen, 1956; Lange. 1959; Desouza, 1966). According to Allen and Allen (1947), Williams (1967) and Lim and Burton (1982), the limited knowledge about the nodulation status of Leguminosae is due to the following major difficulties (i) Inaccessibility of specimens, many of which are found in areas not easily reached by collectors. (ii) Confinement of some genera and species to certain limited parts of the world. (ill) Difficulty of examining woody tree genera for presence or absence of nodules. (iv) Non-availability or short life of seeds of tropical legume species which are required for confirming nodulating ability. (v) The fact that, if not all, most of the promising tropical legumes are 7 poor seed producers, or that the seeds they produce are difficult to harvest for reasons of pod dehiscence, intermediate flowering habit, or buried seed bearing pods in masses of foliage. Beadle (196^) found that out of 80 legumes surveyed in arid or semi-arid regions of eastern Australia, 68 were nodulated. Norris (1969) surveyed rain forest legumes in Amazonia and Guyana and recorded nodules on 30 out of 53 tree species examined. Corby (1971) described the characteristics of wild legumes indigenous to Rhodesia, according to their tribal classification. Gallardo (1970) examined nodulating species in Argentina. Dubey e^_aj_ (1972) surveyed the Island of Puerto Rico for indigenous nodulated legumes. Lim and Ng (1977) examined the nodulation of legumes in Singapore. •They reported for the first time the occurrence of nodules on Adenanthera pavonina. Yanasugondha e t ^ (1977) studied the nodulation of 52 species of native legumes from various parts of Thailand. The nodulating ability of legumes in sub-tropical Pakistan was studied by Athar and Mahmood (1979) who reported 52 species of Papilionoideae as having nodules, five of these being new records. The work done in India on the nodulation of wild legumes spreads over the last three decades - apart from isolated reports like that by Joshi (1920) who worked on root nodule bacteria in Crotalaria juncea. Nodulation on Crotalaria retusa and Clitorea ternatea was shown by Rangaswami and Oblisami (1962). Satyanarayan and Gaur ( 1965) observed that Atylosia scaraboides, Rhyncosia minima and Tephrosia purpurea plants have very deep root systems and they are either devoid of nodules or contain very few, limited to their lateral roots. The nodulation 8 and nitrogen fixation ability of Sesbania cannabina has been studied by Singh (1971). Nodulation in 2^* wild legunne species growing in Hooghley and Burdwan districts of West Bengal was reported by Sinha e ^ ^ (1971). According to Subba Rao (1972) hardly 10% - 12% of leguminous species have been examined for nodulation; and of these, 65% Caesalpinoideae, 10% Mimosoideae and 6% Papilionoideae plants do not bear nodules. Out of the 25 species of tree legumes belonging to the genera Acacia, Albizia, Bauhinia, Colophospermum, Dichrostachys, Leucaena, Peltophorum, Pithecellobium and Prosopis nodulation was observed by Basak and Goyal (1980a) in only 19 species. Nodulation on Crotalaria angulata and Rhyncosia velutina was reported for the first time by Subramanian and Manjula (1986) who also reported that Cassia auriculata and occidentalis lack nodulation although previously reported as nodulating species. Few reports are available on the nodulation of wild legumes in Maharashtra State. Bhelke (1972) reported nodulation for the first time on some species of Alysicarpus monj liferg^ A. tetragonolobus, Clitorj a biflora, Crotalaria filipes, C. nana, Dalbergia sympathetica, Geissaspis cristata, Smithia capitata, S^. purpurea and pycnantha. Nimbalkar . (1986) reported nodulation on 19 legume species during the survey of wild legumes in Western Maharashtra. 2.1.1 Black-coloured nodules. The internal pigmentation in the root nodules of legumes is indicative of their nitrogen-fixing effectiveness. Actively nitrogen-fixing nodules range from pink to red-brown in colour due to the presence of leghaemoglobin. Non­ nitrogen-fixing nodules, on the other hand, are normally white or green in colour. Occasionally black nodules are formed. While screening rhizobial isolates from west Africa for effectiveness in the case of cowpea, Eaglesham et al (1982) detected black-coloured nodules in cowpea. Instances of unusually black-coloured nodules have been reported for several tropical legumes; Vigna mariana (Allen and Allen, 1936), Doiichos lablab (Cloonan, 1963), Mimosa spp. and Leucaena glauca (Campelo and Campelo, 1970), Centrosema pubescens and Phaseolus atropurpureus (Vincent, 1970), Mucuna pruriens (Nimbalkar, 1986). According to Cloonan (1963), the black pigment may be due to melanin formed by the enzyme phenolase in the nodules. Eaglesham e^_a^ (1982) attempted to correlate nodule phenotype with colony morphology and found that only certain dry-colony type strains and not the wet colony-type, caused black nodulation on cowpea. Studies by Raffiquddin (198^) on competition between inoculated and native rhizobia for nodulation of cowpea, Vigna unguiculata Walp. showed the usefulness of black-nodule strain in evaluating nodulating competitiveness of cowpea rhizobia in soils where black-nodule strains are not indigenous. 10 2.2 STUDIES ON ROOT NODULE BACTERIA 2.2.1 Classification : In the 9th edition of Bergey's Manual of Systematic Bacteriology, nodule- forming bacteria have been classified into two genera ; Rhizobium and Bradyrhizobium (Jordan, 198^). Differentiation of these genera is supported to a great extent by cross-inoculation groups. Fast-growing, nodule-forming bacteria having a generation time of less than six hours have been included in the genus Rhizobium whereas the nodule bacteria having a generation time of more than six hours form the genus Bradyrhizobium (Jordan, 1982). In the new system of c la s s ific a tio n , Rhizobium phaseoli and ■Rhizobium trifolii have been fused with Rhizobium leguminosarum which contains three biovars, viz. viceae, trifolii and phaseoli. Rhizobium meliloti (Dangeard) and Rhizobium loti have been designated as separate species (Jarvis 1982). A new species Rhizobium fredii is erected for fast-growing rhizobia that nodulate soybean (Scholia and Elkan, 198^). The genus Bradyrhizobium represents a heterogeneous group of nodule bacteria of which the taxonomic relationships are not well understood. This genus has only one designated species, viz. Bradyrhizobium japonicum. For the present, it is suggested that, members of the genus Bradyrhizobium, other than japonicum, be referred to as Bradyrhizobium sp. with the name of the appro­ priate plant in parenthesis immediately following; e.g. Bradyrhizobium sp. (Vigna). In this classification, no species epithet is given to the members of the cowpea miscellany group. 11 At present not more than approximately 8% of the 1^,000 or so known species of legunninous plants have been examined for nodulation, and less than 0.5% have been studied with respect to their symbiotic relationships with nodule bacteria (Jordan, 1982; Tauro, 1986). Most of the plants examined have been of agricultural importance, and the huge reservoir of wild tropical leguminous plants is only beginning to be investigated. It thus becomes clear that the present classification can not be accepted as final and it is quite likely that it will be gradually modified after a larger number of Rhizobium strains from a wide variety of leguminous plants will be studied. 2.2.2 Cultural and morphological characteristics : The rhizobial cells are aerobic, Gram negative, motile rods measuring G.5 - 3.5 )jm X 0.7 - 3 ym, non-spore forming, capsulated or non-capsulated with polar or peritrichous flagella. The typical rhizobia produce circular, convex, white glistening colonies, with entire margins on Congo red yeast extract mannitol agar
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