that are expressed before the onset of nitrogen fixation and are involved in infection and nodule development. The products of the late nodulin genes are involved in the interaction with the endosymbiont and in the metabolic specialization of the nodule (Nap and Bisseling, 1990). In the first part of this review, we describe the early steps of the interaction between rhizobia and legumes that result in the formation of a nitrogen-fixing nodule. We focus on the role of specific lipooligosaccharides secreted by rhizobia in the induction of these early steps. In the second part, we describe nodule functioning and compare actinorhizal and legume nodules.

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Symbiotic Nitrogen Fixation.

The Rhizobium-plant symbiosis.

Genetically, Rhizobium sp. strain NGR234 and R. fredii USDA257 are closely related. Small differences in their nodulation genes result in NGR234 secreting larger amounts of more diverse lipo-oligosaccharidic Nod factors than USDA257. What effects these differences have on nodulation were analyzed by inoculating 452 species of legumes, representing all three subfamilies of the Leguminosae, as well as the nonlegume Parasponia andersonii, with both strains. The two bacteria nodulated P. andersonii, induced ineffective outgrowths on Delonix regia, and nodulated Chamaecrista fasciculata, a member of the only nodulating genus of the Caesalpinieae tested. Both strains nodulated a range of mimosoid legumes, especially the Australian species of Acacia, and the tribe Ingeae. Highest compatibilities were found with the papilionoid tribes Phaseoleae and Desmodieae. On Vigna spp. (Phaseoleae), both bacteria formed more effective symbioses than rhizobia of the "cowpea" (V. unguiculata) miscellany. USDA257 nodulated an exact subset (79 genera) of the NGR234 hosts (112 genera). If only one of the bacteria formed effective, nitrogen-fixing nodules it was usually NGR234. The only exceptions were with Apios americana, Glycine max, and G. soja. Few correlations can be drawn between Nod-factor substituents and the ability to nodulate specific legumes. Relationships between the ability to nodulate and the origin of the host were not apparent. As both P. andersonii and NGR234 originate from Indonesia/Malaysia/Papua New Guinea, and NGR234's preferred hosts (Desmodiinae/Phaseoleae) are largely Asian, we suggest that broad host range originated in Southeast Asia and spread outward.

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Rhizobium sp. strain NGR234 and R. fredii USDA257 share exceptionally broad, nested host ranges.

Symbiotic mutants of rhizobium meliloti that uncouple plant from bacterial differentiation

A total of 33 strains of fast-growing soybean rhizobia isolated from soil and soybean nodules collected in China and 25 strains belonging to the genera Rhizobium, Bradyrhizobium, and Agrobacterium were compared by numerical taxonomic techniques, using 240 different characters. Our results indicated that all of the strains of fast-growing soybean rhizobia which we examined are closely related (guanine-plus-cytosine content, 59.9 to 63.8 mol%) and are separated from Rhizobium and Bradyrhizobium at the generic level. Based on numerical taxonomy, deoxyribonucleic acid (DNA) base ratio determinations, DNA-DNA hybridization data, serological analysis data, the composition of extracellular gum, bacteriophage typing data, and soluble protein patterns, we propose that the fast-growing soybean rhizobia represent members of a new genus rather than a species of Rhizobium (Rhizobium fredii); we propose Sinorhizobium gen. nov. as an appropriate generic name. The type species of the new genus is Sinorhizobium fredii comb. nov. (basonym, Rhizobium fredii Scholla and Elkan 1984), and the type strain is strain ATCC 35423 (= USDA 205). For the other species we propose the name Sinorhizobium xinjiangensis sp. nov.; the type strain of this species is strain CCBAU 110, which has been deposited in the Beijing Agricultural University Culture Collection, Beijing, People's Republic of China.

Numerical Taxonomic Study of Fast-Growing Soybean Rhizobia and a Proposal that Rhizobium fredii Be Assigned to Sinorhizobium gen. nov.

The infection process in Arachis hypogaea by rhizobia differs from that normally found in Trifolium spp in that no infection threads are formed The root hairs, which are long (up to 4 mm), septate, and often with large basal cells, occur only at the sites of emerging lateral roots Infection occurs only where the root hairs have large basal cells Rhizobia cause curling and deformation of the root hairs (as in Trifolium spp) but enter the root at the junction of the root hair and the epidermal and cortical cells The bacteria are distributed intercellularly via the middle lamellae and enter the cortical cells through the structurally altered cell wall, often close to the host cell nucleus The root hairs and large basal cells become infected in the same way Within the cortical cells of the emerging lateral root the rhizobia multiply rapidly and the invaded cells divide repeatedly to form the nodule tissue Bacteriod formation occurs only when the host cell ceases to divide

Some Observations on Infection of Arachis hypogaea L. by Rhizobium

A low concentration of yeast extract (0·1%) in liquid media favoured rapid growth and high percentage of viable cells in cultures of Rhizobium japonicum (CB 1809), R. lupini (WU 425), R. meliloti (SU 47), R. trifolii (TA1) and a cowpea strain (CB 756). Concentrations of yeast extract > 0·35% depressed viability and produced distorted cells in all strains except SU 47: TA1 was especially sensitive. When used at 0·5–1% (w/v), each yeast extract (Difco, Oxoid, Vegemite) or casein hydrolysate produced greatly enlarged abnormal cells of TA1, each containing several granules of poly-β-hydroxybutyrate and whorls of intracytoplasmic membranes, and showing greater internal disorganisation than that seen in root nodule bacteroids. Lysogenic and non-lysogenic cultures of R. trifolii were all sensitive to yeast extract, and such sensitivity, for strains of several species, was unrelated to effectiveness in nodulating host plants. Glycine inhibited growth of all strains tested. Several other amino acids occurring in casein hydrolysate inhibited TA1 strongly and induced formation of distorted cells and spheroplasts; this distortion was partly counteracted by adding salts of calcium or magnesium. In media with 0·1% yeast extract the use of mannitol, sucrose, lactose or galactose as alternative carbon sources, each at a concentration of 0·02–1%, did not affect numbers of viable rhizobia or cell shape in all strains tested.

Effect of Yeast Extract Concentration on Viability and Cell Distortion in Rhizobium spp.

The Rhizobium leguminosarum host range plasmid pJB5JI was transferred to three fast-growing bacterial strains able to nodulate soybeans. These strains, isolated in China, contained plasmids and were able to transfer pJB5JI back to R. leguminosarum. Soybean strains carrying pJB5JI elicited early stages of nodule development on peas.

Muriel R. Chandler

Papers

Some Observations on Infection of Arachis hypogaea L. by Rhizobium

Effect of Yeast Extract Concentration on Viability and Cell Distortion in Rhizobium spp.

Transfer of a host range plasmid from Rhizobium leguminosaum to fast‐growing bacteria that nodulate soybeans