wild and weed azuki beans in japan

WILD AND WEED AZUKI BEANS IN JAPAN 1

HIROFUMI YAMAGUCHI

Yamaguchi, Hirofumi (College of Agriculture, University of Osaka Prefecture, SakaL Osaka, 593 Japan). WILD AND WEED Azurd BEANS IN JAPAN. Economic Botany 46(4):384-394. 1992. WiM azuki bean, a progenitor of an Asiatic food legume (Vigna angularis var. nipponensis: Fabaceae), and its weed form are distributed widely in the Japanese Archipelago. The straggling or climbing wiM form occurs in sleeve or mantle plant communities, and the weakly climbing or bushy weed form is found in relatively open human-disturbed habitats. The wiM form has small seeds with a black-mottled pattern on green or grey skin; the weed form has larger seeds with variable color patterns. WiM and weed forms have black, easily dehiscent pods, distinct from their cultivated counterpart which has red large seeds and indehiscent light-colored pods. The wild form is not utilized, but the weed form is recognized by farmers and has several folk names as a weed, a contaminated form of azuki bean, and a substitute for azuki as a food. The frequent occurrence of weed azuki bean in Japan is attributable to adaptation of the wiM form to lack of climbing support in human-disturbed habitats, escape from old cultivars, and natural establish- ment from the derivatives of hybrids between cultivars and wild forms.

Wild- und Unkrautformen der Azuki Bohne in Japan. Die Wildazukibohne, ein Vorfahr einer asiatische Eflbarbohne (Vigna angularis vat. nipponensis: Fabaceae), und dieser Unkrautform sind weitverbreit in der japanischen Inselgruppe. Die iippigwachende oder aufsteigende Wildform liegt in den Saum- oder Mantelpflanzengesellshaften, und die schwiichliche aufsteigende oder buschige Unkrautform wird in den relativoffen menschenausst6rend Standorten gefunden. Die Wildform trage die kleinen Samen mit einem schwarzgefleckten Muster auf der grauen oder griinen Samenhaut; die Unkrautform trage die grb'fleren Samen mit dem variabelen Fiirbung und Zeichnung. Wild- und Unkrautformen haben die schwarzen und lechtspaltigen Hiilsen, distinktem vom ihnen Kulturgegenstiick, der hat die roten groflen Samen und die inspaltigen hellfarben Hiilsen. Die Wildform wird nicht gebenutzen, abet die Unkrautform wird angeerken- nen bei den Bauern und hat fieln Volksnamen als einen Unkrauten, einen Befleckung/'ormen der Azukibohne, und zum Ersatz J~r Azuki zu Efizwecken. Das hiiufige Dasein der Unkrautazuki- bohne in Japan ist der Anpassung der Wildform zu Mangel der aufsteigenden Stiize in den menschenaussfrenden Standorten, dem Verwilderung yon alten Kulturvarietiiten, und der na- tiirlichen Griindung yon der Ableitung der Hybriden zwischen Kulturvarietiiten und Wildformen hervorgekom men.

Key Words: azuki beans; weed azuki beans; wild azuki beans.

There is continued interest in how our cultivated plants have been domesticated from their wild progenitors. Domestication is the process of accumulation of plant-human interactions (Rindos 1984) that increase morphological, physiological and ecological changes in wild species and enable them to adapt to conditions under human cultivation (Harlan 1975). Compar- ative analyses of the morphology of domesticates and their wild counterparts illustrate distinct changes in gigantism and several features on dis-

Received 3 February 1992; accepted 4 August 1992.

persal and self-growing abilities (Harlan 1965; Heiser 1988; Nakao 1976; Schwanitz 1966). The occurrence of weed races among cultivated plants in nature has played an immense role in our understanding of the process of domestication (De Wet and Harlan 1975; Harlan 1965; Vavilov 1926), though delimitation of weed and wild races is very difficult because of our limited knowl- edge of wild relatives of cultivated plants.

Although grain legumes have been among the most important human food crops and along with the cereal grain crops have had the strongest con- nections to the evolution of civilization, the domestication of Asiatic food legumes remains

Economic Botany 46(4) pp. 384-394. 1992 �9 1992, by The New York Botanical Garden, Bronx, NY 10458 U.S.A.

1992] YAMAGUCHI." AZUKI BEANS 385

problematic (Smartt 1990). The azuki bean ( Vig- na angularis (WiUd.) Ohwi et Ohashi: subgenus Ceratotropis) is a minor but very important crop; its red bean is used in dishes eaten to commem- orate formal occasions like births, weddings, deaths, and private and tribal festivals in East Asia. Var. nipponensis (Ohwi) Ohwi et Ohashi, which is considered to be the wild counterpart of var. angularis (Marrchal et al. 1978; Ohwi and Ohashi 1969; Tateishi and Ohashi 1990), is distributed on the main islands of Japan (Hon- shu, Shikoku and Kyushu) and in Korea, Tai- wan, southeastern China and Nepal (Ohwi 1953; Tateishi 1983, 1984). Recently it was found that a morphologically intermediate form between cultigen and true wild nipponensis grows natu- rally and that farmers recognize the form as a "weed" (Yamaguchi 1989). These circumstances enable us to better understand the process of domestication. On the basis of my exploration of wild azuki-bean populations throughout Ja- pan, I describe features of plant architecture, pod and seed, and distribution of wild and weed azuki beans in Japan, and discuss the interaction among wild, weed and domesticated races.

METHODS

Explorations were made in Hokkaido, Hon- shu, Shikoku, Kyushu and the Ryukyus during the period from late flowering to maturing (Sep- tember to October) in 1988, 1989, 1990 and 1991. In every site, associated dominant plant species and climbing support were recorded. Fully ma- ture pods just prior to ejecting seeds were gathered from every population. The features of pods and seeds were recorded, and their sizes were measured. Pods of cultigens were collected from farmers' fields neighboring the wild populations in order to compare general morphology. A small number of seeds scratched by sand paper were sown to investigate juvenile growth. To clarify the limitation of distribution, a herbarium sur- vey was conducted at TNS, TI, TUS, MAK and KYO herbaria.

RESULTS

HABITAT AND GROWTH FORM OF WILD AND WEED AZUKI BEANS

Natural azuki populations were found in mantle plant communities at sloping moist sites and in human-disturbed mesic sites, and their plant stature shows clear discontinuity between pop-

ulations from different habitats in Japan. In natural populations in mantle and sleeve plant communities, 15 to 50 individuals were crowded. This representative is considered to be a true wild form, and its thin twisted branching stems usually climb to the branches and stems of Artemisia princeps Pamp., Miscanthus sinensis Anderss., Reynoutria japonica Houtt., Bidens biternata Merr. et Scheff, Achyranthes bidentata Blume var. tomentosa Sieb. et Zucc., Solidago altissima L., Sonchus oleraceus L., Erigeron canadensis L., Helianthus tuberosus L., Pleioblastus simonii Nakai, Ambrosia trifida L., Xanthium occiden- tale Bertoloni (Fig. 1, 2). The wild form grows together with some associated herbaceous and vine species, Commelina communis L., Setaria viridis Beauv., Kalimeris yomena Kitam., Per- sicaria longiseta Kitag., Rubus hirsutus Thunb., Amphicarpaea bracteata Fernald. subsp, edge- wortii Ohashi var. japonica Ohashi, Humulus ja- ponicus Sieb. et Zucc., Glycine max Merr. subsp. soja Ohashi, Cayratia japonica Gagn., Pueraria lobata Ohwi. In human-disturbed habitats, relatively small numbers of individuals grow; their erect thick stems grow with some annual weed (Fig. 3), and they frequently infest the upland fields planted in soybeans (Glycine max), azuki beans, and tea plants (Cammelia sinensis L.). This representative, considered to be a weed form, infrequently elongates the tip of its loosely twisting central stem that leans on the crops, the leaves and stems of Artemisia princeps, Miscanthus sinensis, Imperata cylindrica Beauv., Pleioblastus simonii, planted trees, artificial poles and nets. Thus, among natural azuki populations in human-disturbed habitats, especially in weedy places, the weed form is easily distinguishable from the "true" wild form by its morphological and ecological features, which are intermediate between wild and cultivated azuki beans.

GENERAL FEATURES OF WILD, WEED AND CULTIVATED AZUKI BEANS

Wild, weed (intermediate) and cultivated forms are strictly annual: their seeds germinate in early summer, they bear flowers in late summer, and their seeds bear fruit in the autumn. Three forms share conspecific taxonomical characters (stip- ule, bracteole, pistil, and hilum shape), and morphological differences among the three forms are distinct particularly in plant architecture, dehiscence and color of pods, and color and size of seeds (Fig. 5, 6). In the wild azuki, all the plants

386 ECONOMIC BOTANY [VOL. 46

Fig. 1-4. Fig. 1. Wild azuki in bloom in a relatively open habitat. Fig. 2. Wild azuki twisting around the stem of Reynourtria japonica Houtt. Fig. 3. Weed azuki in a kitchen garden abandoned for a few years. Fig. 4. Domesticated azuki bean in a swidden in Ishikawa Prefecture.

1992] YAMAGUCHI: AZUKI BEANS 387

Fig. 5. Morphological explanation of the wild azuki bean. A, whole plant (• B, inflorescence (x 1.5); C, flower ( x 2); D, standard vexillum ( x 2); E, wing ( x 2); F, keel ( x 2); G, flower without corolla ( x 2); H, calyx and pistil ( • 3); I, calyx and bracteoles ( x 3); J, legume ( x l); K, single seed ( x 3).


Fig. 6. Morphological explanation of the weed azuki bean in comparison with Vigna angularis var. angularis (domesticate). A-F, weed form; G-K, cultivated form; A, whole plant (x0.5); B and G, inflorescence (x 1.5); C and E, flower (x 2); D and I, calyx and bracteoles (x 3); E and H, legume (x 1); F and K, single seed (x 3).


Fig. 7. Seedlings of wild (right), weed (center) and cultivated (left) azuki beans. Bar indicates 5 cm.

show slender plant stature and rapid dehiscence of black pods (5.5-7.2 cm in length); the racemous inflorescence with small flowers usually towers above the canopy (Fig. 1), and pod clusters are located distant from the rooting place as a result of the plants ' straggling and climbing architecture. Though the plant shown in Fig. 5 shows a lack of lateral branching at the lower axils, the wild form usually sends out 2--4 lateral branches with many branchlets over 1 m long. In contrast, the weed form features bushy or slightly climbing plants without any lateral branching at the lower axils. Its pod clusters, with easily dehiscent black pods (6.1-8.2 cm in length), cling behind the leaf canopy in open habitats (Fig. 3), and frequently the weed form shows towering racemous flowers in the mantle vegetation and elongated and twisted stems on the upper part of the plant. On the other hand, the domest icated azuki bean has large hard-dehiscent colorless or straw-colored pods (7.5-15 cm in length). Its inflorescence with large flowers does not display over the canopy (Fig. 4), and pod clusters are situated near the main stem. No sig- nificant difference among three forms was found in the number of seeds per pod. Pod wall of the cultivated form is papyriferous and monil i form but that of wild and weed forms is striated co- riaceous.

In natural populat ions the height of first leaves fluctuates with environmental condit ions as well

as with the posit ion of branching, then obser- vation of seedlings gives a clear answer whether a plant is a weed or wild form. A clear discon- t inuity between wild and weed forms is found in the height of first opposite simple leaves (Fig. 7) and first long pr imary branches. The wild form expands its first leaves from the dark purple main stem and extends lateral branches horizontally in close contact with the soil from axils of the first leaves and subsequent cauline leaves. The weed form expands its first leaves from the green main stem ca. 6-10 cm above ground level and extends its first long branch from the fourth or fifth node, as is seen in the cultivated form. Seed- lings of cult ivated form show vigorous growth, with large leaves and a thick green stem. Pr imary leaves respond to temperature fluctuation im- mediately in the wild forms, moderately in the weed forms and sluggishly in the cultivated forms.

Average sizes (length x width x th ickness)of seeds range from 3.65 m m x 2.76 m m x 2.37 m m to 4.59 m m x 3.69 m m x 3.30 m m in wild forms, from 4.26 m m x 3.24 m m x 2.79 m m to 5.78 m m x 4.55 m m x 4.30 m m in weed forms, and from 5.20 m m x 3.54 m m x 3.03 m m to 8.39 m m x 6.17 m m x 5.76 m m i n cultivated forms. Wild forms all show a black mott led pattern on the grey or green skin of the seed coat. Weed forms show variable seed colors: yellow brown, grey, yellow green and brown mottled patterns on the light brown skin, or the same


patterns as in the wild. In contrast, the cultivar usually has a red skin, and frequently is white, black, black mottled on red skin, variegated red and white, or very rarely the same seed coat types as in the weed form.

GEOGRAPHICAL DISTRIBUTION AND SEED COAT VARIABILITY OF WILD AND

WEEDY POPULATIONS

Although herbarium specimens tell us little about growth habits and seed colors, the holotype ofvar , nipponensis (Ohwi, J. s.n. KYO) shows a wild type of plant architecture. Specimens ofvar. nipponensis deposited in these herbaria were collected from the Honshu, Shikoku and Kyushu islands of Japan, Korea, Taiwan, China and Ne- pal, and representatives from eight localities in the specimens deposited in various herbaria (To- gashL M. s.n. TI; Tateishi 679 TI; Tateishi 483 TUS; Ohashi, Tateishi & Ohba 169 TUS; Hatsu- shima, Seko and Kanno s.n. MAK; Araki 13795 KYO; Horie, K. s.n. KYO; Kurosaki 13701 KYO) are assumed to be weed forms based on their pod and seed features (Fig. 8). On the basis of my collections, it appears that wild and weed forms are distributed widely in Japan except for Hok- kaido and the Ryukyu islands. In the seed color of wild and weed forms, the phenotype of black mottled pattern on grey or green skin is distributed throughout Japan, but other phenotypes show restricted distribution. In the weed form, the phenotypes with pure brown and brown mottled on brown skin are distributed in the Kinki district, the grey phenotype in southern Honshu and Kyushu, and the green phenotype in Kyushu. There is no red colored seed in the 94 natural populations investigated, except in one putative hybrid population.

Most of the individual populations showed monomorphic narrow variation in size and color of the seed and pod (Fig. 8); five populations showed dimorphic discontinuous variation, and only one population showed a wide continuous variation from large red seeds to small black mottled seeds. The former five populations seem to be characterized by coexistence of wild and weed forms, because a few plants in these populations have larger pods, larger seeds and short- er plant stature. The unique population with wide variation, located on the Boso peninsula in Chiba Prefecture, seems to be a hybrid swarm between cultivated and wild azuki beans. In this population all individuals were climbers twisted

around Miscanthus sinensis at the slope edge of an upland field planted in cultivated azuki beans. The variation pattern for seed and pod traits distinctly shows that this population may be composed of hybrid derivatives and the mem- bers ofvar, nipponensis, because it contains various combinations of cultivated-type and wild- type pods with a black mottled pattern and brown, grey and red seeds. The variability of this putative hybrid population will be described else- where in detail. No other natural population shows red seeds and indehiscent colorless pods.

UTILIZATION AND FARMERS' RECOGNITION OF WILD

AND WEED AZUKI

Although the wild form is never utilized or recognized as food or other plant resource in Ja- pan, the weed form of azuki is utilized as a substitute for the cultivated form and is recognized as an azuki-l ike bean called " N o r a - a z u k i , " "Nouraku-azuk i , " "No-azuk i , " " N o r a k k o , " "Yama-azuki ," "Kuro-azuki ," "Kitsune-azuki," "Taito-azuki," "Ishimame," "Bundo," and "Masara." The words "Nora," "Nouraku," "No" and "Norakko" in Japanese mean "o f the field" or"ofagricultural sites." The seeds of weed azuki are consumed as the sweet beans "An" or "Anko" and not used in "Azukigayu" (the rice gruel mixed with azuki beans) or "Okowa" (the glutinous rice steamed with red beans) in Japan. In a lean year or when pests attack azuki beans, farmers collect the seeds from the weed form to consume im- mediately and to secure a complement of azuki cultivars for the following year. In some places, the seeds of weed forms are used in beanbags. When the weed form infests cultivated gardens, however, it is recognized as a contamination of azuki beans that lowers seed quality of azuki cultivars. Thus the weed form is preferred to the wild form by people.

DISCUSSION

Morphological and ecological differences between wild and cultivated azuki beans are distinct (Fig. 1-7), as was recognized by Yamaguchi (1989), and they are in many grain legumes (Evans 1976; Gentry 1969; Smartt 1990). In growth habits, plant type, branching pattern, fruiting habits, pod and seed size, and seed coat variability, weed azuki shows an intergradation between wild and cultivated azuki beans. Weed azuki has a plant


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Fig. 8. Geographical distribution of wild and weed azuki beans. A, herbarium record. Filled circle, Vigna angularis var. nipponensis; Open circle, putative weed form. B, author's confirmation. Open circle, weed form; filled circle, wild form; filled circle with a hair, mixed population with wild and weed forms; triangle, putative hybrid swarm.


architecture similar to that ofcult igen and a seed- dispersal and pod dehiscence habit similar to that o f the wild form. These circumstances may give rise to diverse hypotheses on the origin of the weed races: they are variously thought to be (1) exact progenitors of the cultigen, (2) escapees from old cultivars and (3) derivatives from a hybrid between wild and cultivated races (De Wet and Harlan 1975). The view that wild azuki, var. nipponensis, has straggling or climbing growth habits has come to be easily accepted since the known wild Vigna species belonging to subgenus Ceratotropis in Japan, 1I. nakashimae Ohwi et Ohashi, 1I. reflexo-pilosa Hayata and 1I. minimus Ohwi et Ohashi var. minor Tateishi, show straggling, lean or climbing growth habits as vines associated with mantle and sleeve plant communit ies that are mainly composed of Mis- canthus sinensis and Artemisia princeps. In contrast, the bushy plant architecture of weed azuki is considered to be an advantage in human-disturbed habitats where no climbing support is available, though the weed form is able to grow even in mantle and sleeve sites because of its high phenotypic plasticity. The change of plant form may be essential for the evolution of domestication in the azuki bean as a hand-gathered grain legume irrespective of the evolution of lack of seed dispersal abili ty and lack o f seed dor- mancy, which are assumed to have evolved under cult ivation (Blumler 1991; Ladizinsky 1987). Thus, weed azuki may be a progenitor of the cultigen.

The second hypothesis is also supported by the evidence that the traits specifying wild and weed forms are found independently in some azuki cultivars. The off-type or weed form could easily be the result of a recombinat ion or the addi t ion of new mutations. Cultivars similar to the weed form, which have relatively easily de- hiscing pods containing seeds with a black mottled pattern on grey skin, had been cult ivated as "Kage-azuki" in Japanese swidden (Noda 1952). The folk name of the weed race, "Noraku-azuki," appeared as a local cultivar in an old Jap- anese agricultural manual, Shika Nougyou Dan (Miyanaga 1789). This evidence clearly suggests past uti l ization of the weed form and occurrence of pr imit ive cultivars. The quality of the azuki bean may have been mainta ined by careful ex- aminat ion by the farmers and by rotational crop- ping that excluded off-types and weed forms from

inclusion among local varieties. Off-types or old cultivars could be grown under human-dis turbed habitats that were still wild.

The last hypothesis, that of hybrid origin, is the most probable in the form of contaminat ion of small mott led seeds into azuki bean (Egawa et al. 1990). However, lower variabil i ty in wild and weed populations, except in one putat ive hybrid population, does not suggest that hybridizing is the only factor. Since most of the cultivated azuki in Japan are red-seeded types that are controlled by 7 major genes, RRhhgg- f fmmccZZ (Takahashi and Fukuyama 1917), and most wild forms have black mott led seeds with genes RRHHGGffAIMccZZ, i f the weed form were a derivative of a hybrid between wild and cultivated forms with subsequent self-pol- linations, an even occurrence of red-seeded and various colored phenotypes would be expected. The lack of a red-seeded phenotype in the weed form is interpretable only if there is differential selection against red or non-red. Since domestication is a process of accumulat ion of cultivated features and may include hybridizing and diver- sifying between wild and cultivated counterparts (Harlan 1965), the weed form may include tran- sitions from wild forms to cultigens, escapes from old cultivars and derivatives from hybrid between wild and cultigen.

Regardless of how the weed form originated, the extensive and wider distr ibution of the weed form in Japan and the existence of a hybrid swarm between wild and cultivated azuki beans clearly supported the assumption that var. nipponensis is a progenitor of var. angularis which has been postulated due to close morphological similari- ties (Marrchal et al. 1978, Tateishi pers. comm.). The assumption is also supported by the same chromosome number (Sawa 1983) and high cross- compatibi l i ty (Sawa 1983; Siriwardhane et al. 1991), a similar RFLP pattern in cp D N A (Mi- kami et al. 1991) and a similar SDS page profile of seed-storage proteins (Yamaguchi and Kosuge 1991) between them. In contrast, there is distinct differentiation in gross morphological features between races that relate closely with regard to habitat. These features clearly suggest that the divergence of the azuki bean into three races may be a relatively recent event, and thus that domestication of azuki bean is a process of infraspecific differentiation of some ecologic and ag- ronomic characters.


I have no ideas on the botanical posi t ioning o f the weed race under which open or closed systems should be appl ied (Brandenburg 1986), because the weed race m a y be an aggregate o f t ransi t ions f rom wild to domest ic , escapes f rom old cul t ivars a n d / o r der iva t ives f rom a hybr id between cul t ivar and wild races. Since wild var. nipponensis is d is t r ibuted in the warm tempera te zone that includes Nepal , cont inenta l China, and Taiwan (Ohashi et al. 1988; Tateishi 1983, 1984), and since in teract ion a m o n g wild types and cultigens is not fully known in areas outs ide Japan, it is not possible to say more about the origin o f the domes t ica ted azuki at this t ime.

A C K N O W L E D G M E N T S

I gratefully acknowledge support from the following: the Japan Beans and Peas Foundation for partial financial support; M. Umebayashi for fine illustrations; Y. Ishimine, M Kataoka, K. Nishibori, T. Ogata, H. Shibayama, Y. Tateishi, K. Terai, K. Watanabe and S. Yamaguchi for assistance with my field surveys; and S. Moriuchi for the German trans- lation.

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Evans, A . M . 1976. Beans. Pages 168-172 in N. W. Simmonds, ed., Evolution of crop plants. Long- man, London.

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- - , and J . M. J. de Wet. 1965. Some thought about weeds. Econ. Bot. 19:16-24.

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Marrehal, R., J. M. iaseheroa, and F. Stainier. 1978. Etude taxonomique d'un groupe complexe drsprcies des genres Phaseolus et Vigna (Papilion- aceae) sur la base de donres morphologiques et pol- liniques traitres par ranalyse informatique. Bois- siera 28:1-273.

Mikami, T., S. Kato, H. Yamaguehi, and Y. Shima- moto. 1991. Chloroplast DNA diversity in azuki bean and its relatives. Jpn. Jour. Breed. 41, Supple. 1:476-477.

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(V. angularis) by interspecific crossing. Michurin Biol. Res. 18:3-26.

Schwanitz, F. 1966. The origin of cultivated plants. Harvard Univ. Press, Cambridge, MA.

Siriwardhane, D., Y. Egawa, and N. Tomooka. 1991. Cross-compatibility of cultivated adzuki bean ( Vig- na angularis) and rice bean (V. umbellata) with their wild relatives. Plant Breeding 107:320-325.

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Takahashi, Y., and J. Fukuyama. 1917. Morpholog- ical and genetic studies on the adzuki bean. Rep. Hokkaido Agric. Exp. Sta. 7:1-185.

Tateishi, Y. 1983. Leguminosae collected in the Arun valley, East Nepal. Pages 134-146 in M. Numata, ed., Structure and dynamics of vegetation in east- ern Nepal. Chiba University, Chiba.

�9 1984. Contribution to the genus Vigna (Le- guminosae) in Taiwan I. Sci. Rep. Tohoku Univ. 4th ser. (Biol.) 38:335-350.

, and H. Ohashi. 1990. Systematics of the azuki bean group in the genus Vigna. Pages I89-199 in K. Fujii et al., eds., Bruchids and legumes: eco- nomics, ecology and coevolution. Kluwer Publ., Netherlands.

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, and K. Kosuge. 1991. Evaluation of wild azu-

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A N N O U N C E M E N T

J. Travis Columbus at the Universi ty Herbar- ium, Universi ty of California at Berkeley, is the recipient of the 1992 Lawrence Memorial Award. A student of Dr. Thomas Duncan, Mr. Colum- bus has undertaken a study of the generic-level taxonomy and evolution in the grama grasses and relatives (Gramineae: Cynodonteae: Bou- telouinae). The proceeds of the Award will help support his field research, especially in Mexico.

Commemora t ing Dr. George H. M. Lawrence, founding Director o f the Hunt Institute for Bo- tanical Documentat ion at Carnegie Mellon Uni- versity, the semi-annual Award of$1000 is made to an outstanding doctoral candidate for travel in support of dissertation research in systematic botany or horticulture, or the history of the plant sciences.

wild and weed azuki beans in japan

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