Review Paper The Origin of Cultivated Buckwheat in Mankang District of the Sanjiang Area of Eastern Tibet and its Diffusion to India and the Himalayan Hills Ohmi OHNISHI Kyoto University, Kyoto, Japan DOI https://doi.org/10.3986/fag0042 To the memory of Ohmi Ohnishi, Professor Emeritus of Kyoto University and Editor Emeritus of FAGOPYRUM journal is re- printed the paper, originally published in FOLIA BIOLOGICA ET GEOLOGICA 61/1 (2020), 7-15. Key words: buckwheat, wild ancestor, origin, diffusion, India, Himalaya, short day plant ABSTRACT Natural populations of the wild ancestor of cultivated common buckwheat were searched and collected, starting from its discovery in1990 and finishing the collections in 2005. Among the collections, the samples Zhuka, Xihe from Mankang district of Tibet are most closely related to cultivated common buckwheat. On the other hand, cultivated populations of common buckwheat in Zhouba, Zhubalong both from Mankang district are most closely related with the wild ancestor of common buckwheat. This leads to the hypothesis on the origin of cultivated buckwheat in Mankang district in the Sanjiang area. The diffusion rout from the original birthplace to India and the Himalayan hills is proposed. Several characteristics of Indian and Himalayan common buckwheat are discussed. A main conclusion of the discussion is that the European buckwheat is not of Indian origin nor of the Himalayan origin. It probably came from the northern China through the Silk Road 57 Fagopyrum 41 (2): 57-63 (2024) 1. INTRODUCTION Since the time of de Candolle (1883), (a) what is the wild ancestor of cultivated common buckwheat and (b) where is the original birthplace of cultivated buckwheat, these two problems have been the main issues to be solved by buckwheat scientists. The wild ancestor of cultivated buckwheat was clar- ified as F. esculentum ssp. ancestrale Ohnishi, which was first discovered in 1990 in the Wulang river valley in Yongsheng district of Yunnan province, in China by Ohnishi (1991). As for the birthplace of buckwheat, Ohnishi (2004, 2007, 2010, 2016, and 2018) has repeat- edly claimed that the Sanjiang area of Yunnan, Sichuang provinces and east Tibet in China is the original birth- place of cultivated buckwheat. In the Himalayan countries, India, Nepal, Bhutan, and Pakistan, buckwheat is cultivated extensively and buckwheat is consumed well. Buckwheat in India and the Himalayan hills has some characteristics which are not seen in other regions such as China, Japan and European countries. Today, I discuss the original birthplace of buckwheat in more details, and I consider the diffusion rout from the place of origin to India and to the Himalayan hills. Finally I discuss the characteristics of Indian and the Himalayan buckwheat. As a conclusion, I suggest that the cultivated buckwheat in European countries has never come from India nor from the Himalaya, it probably came from northern China through the Silk Road. 2. THE EXACT ORIGINAL BIRTHPLACE OF COMMON BUCKWHEAT The wild ancestor of cultivated common buckwheat was first discovered in Yongsheng district in Yunnan province of China in 1990 (Ohnishi, 1991). Ten-years searches for the wild ancestor in Yunnan and Sichuan provinces, and the searches in Mankang district of east- ern Tibet in 2002 and 2004, and finally the searches in the Tongyi river valley and the Nyiru river valley in 2004 and 2005 clarified the distribution areas of the wild an- cestor of common buckwheat (Ohnishi, 2007, see also Ohnishi and Tomoyoshi, 2005). The wild ancestor is distributed in 1. The Sanjiang area of Yunnan province and eastern Tibet, 2. Sporadic distribution in northwestern Yunnan prov- ince and southwest corner of Sichuan province, and 3. The Tongyi river valley in Muli district of Sichuan province and the Nyiru river valley in Shangrila district of Yunnan province. Both the Tongyi river and the Nyiru river are small tributary of the Shui- luo river, a tributary of the Jinshajiang river. Among the collected wild ancestor populations, the wild ancestors from the Sanjiang area were revealed to be the most closely related with cultivated common buckwheat in AFLP variation (Konishi et al., 2005) and allozyme variability (see Figure 1, see Ohnishi and Nishi- Photo 1: Cultivated common buckwheat in Sanjian area (Weixi district, Yunnan provice). Flower color in this area near the original birthplace is beautiful pink. Photo 2: The wild ancestors growing in the Xihe river valley of Mankang district in eastern Tibet. The wild ancestors in this valley are genetically most closely related with cultivated buckwheat. Hence, the Xihe river valley along with the towns in northern Man- kang district are considered as the original birthplace of common buckwheat. 58 Ohnishi (2024): The Origin of Cultivated Buckwheat moto, 1988 for the procedures of the electrophoresis and the names of enzymes analyzed). A part of the data on the frequencies of allozymes can be found in Ohnishi (2007). The N-J tree of Figure 1 was written following by Saitou and Nei (1987) using PAUP* version 4.0 (Swof- ford, 1990, 2002). The wild ancestors in the Tongyi river valley and the Nyiru river valley are highly variable in AFLP and al- lozymes, however, the populations from those valleys are most distantly related with cultivated populations of common buckwheat (Fig. 1). The sporadically distributed ancestral populations in Yunnan and Sichuan provinces showed intermediate closeness to cultivated buckwheat (see Fig. 1). This may leads to the conclusion that the Sanjiang area is the original birthplace of common buckwheat. Konishi and Ohnishi (2007) showed that close genetic relation between the wild ancestor in the Sanjiang area and cultivated populations is not due to recent hybridiza- tion between them. Figure 1. The N-J tree among the populations of wild ancestor and cultivated populations of buckwheat based on allozyme data. The name of locations written in Chinese letters in the figure should be expressed in pinyin as is shown in blackets. Only the loctions appeared in text are listed bellow. 西河 (Xihe) 竹卡 (Zheke) 阿東 (Adong) 塩井 (Yanjing) 博科 (Boke) 塩源 (Yanyuang) 金安 (Jinan) 永勝 (Yongsheng) 尼通 (Nidong) 東義 (Tongyi) 依吉 (Yiqi) 竹巴龍 (Zhebalong) 巴塘 (Batang) 肋巴 (Zhouba) N-J tree based on allozyme data Cultivated populations Dongyi river Valley populations Sanjian area populations Photo 3: The cultivation of common buckwheat in Yanjing town of southern Mankang district. The brown part of the cultivated field is common buckwheat just before harvest. Yanjing town has a good weather condition for buckwheat cultivation and the wild ancestors of commom buckwheat are also growing at the margine of buck- wheat fields, although those wild ancestors are not so closely related with cultivated buckwheat. Photo 4: The landscape of northern Mankang district. Although this area is believed to be the original birthplace of common buck- wheat, barley is mainly cultivated in cultivation fields. 59 Fagopyrum 41 (2): 57-63 (2024) Now, by observing Fig. 1 more carefully, you may find that the Zhuka population and the Xihe population from Mankang district are most closely related with cultivated populations. The Adong population (in Yunnan province) and the Yanjing population (close to the border between Yunnan and Tibet, see Fig. 2) are both from the Sanjiang area, but they are slightly far away from the cultivated populations as compared with the Zhuka and the Xihe popultions (Fig. 1). Furthermore, by observing Fig. 1 from the cultivated population side, you may find that the cultivated popula- tions of Zhubalong and Zhouba (both come from north- ern Mankang district) are closely related with the wild ancestor of cultivated buckwheat (see Figs. 1 and 2). Now, as a conclusion, we can say that the Yunnan part of the Sanjiang area is not involved in the origin of buck- wheat cultivation, rather, Mankang district of the San- jiang area, particularly, north part of Mankang district is the original birthplace of cultivated buckwheat. So, it is reasonable that Wang (1986) re- ported archaeological remains (buckwheat seed grains) in the archaeological site of Karuo village near Chamdu of east Tibet. This site is close to the original birthplace Mankang district of eastern Tibet (see Fig. 3). 3. DIFFUSION OF CULTIVATED BUCKWHEAT TO INDIA AND THE HIMALAYAN HILLS Cultivated common buckwheat migrated from its original birthplace, the Sanjiang area of southwestern China to northern China first, then to the Korean pen- insula and Japanese islands (Murai and Ohnishi, 1996). From northern China, cultivated buckwheat migrated west, to the central Asian countries, then to European countries through the Silk Road as I discussed in the pre- vious 13th ISB (Ohnishi, 2016). From the original birthplace of common buckwheat, Mankang district, cultivated buckwheat first went west (Murai and Ohnishi, 1996), overcoming high moun- tains, the Hengdan mountains, in the three- river region (the Sanjiang area), and entered to the Yaruzanpu river basin, then finally arrived at Bhutan, Sikkim, Nepal and India. There exists only one rout connecting Mankang district in the Sanjian area and the Yaruzampu basin in Tibet as seen in Fig. 3. After arriving to these countries, it is easy for buckwheat to travel further west along the Himalayan hills, because the Himalayan hills may pro- vide a comfortable cultivated conditions for cultivated buckwheat. Only a few crops diffused through the same rout as buckwheat. Tea plant, Camelia chinensis, originated in Yunnan province in China, diffused through the same rout as common buckwheat and became an important cultivated plant in India and the Himalayan hills. Only the crop diffused opposite direction from the Himalayan hills to the Sanjiang area is the finger millet (Eleusine cora- cana), originated in Africa. It arrived at the Indian sub- continent, India and Pakistan. Then it diffused to south- ern China, through the same rout as of buckwheat, but in the opposite direction (Hoshikawa, 1992). As shown in Fig. 3, the Karuo archaeological site, from where the oldest buckwheat seed grains were reported, is located not so far away from the original birthplace of cultivated buckwheat. Along the diffusion rout, wild per- ennial buckwheat, F. cymosum ssp. pillus (syn. Fagopyrum Figure 2. Distribution of the wild ancestor of cultivated common buckwheat in Mankang district of the Sanjiang area. : village or town where the wild ancestor was found. Northern population such as Zhouba, Zhuka and Xihe are close to cultivated populations (see Fig. 1) 60 Ohnishi (2024): The Origin of Cultivated Buckwheat pillus Chen, see Chen, 1999) and a weed species F. gra- cilipes are found, near to Dongmai village, Bomi district of Tibet, and Paro of Bhutan, respectively (see Fig. 3). It is well-known that F. cymosum growing in the west of the Yaruzampu grand canyon is all tetraploid, and is of- ten called F. dibotris in Nepal and India (see Hara, 1972). 4. CHARACTERISTICS OF BUCKWHEAT IN INDIA AND IN THE HIMALAYAN HILLS Here, I mention several characteristics of buckwheat and buckwheat cultivation in India and the Himalayan hills. 4-1. buckwheat cultivation as a fresh vegetable in India Both common buckwheat and Tartary buckwheat are cultivated as a fresh vegetable in India, rather than as a grain crop. As a result of long history of cultivation as a fresh vegetable, local varieties for such purpose have become the varieties with very small grains. I observed such a small grain variety in Bageshwar town, the state of Uttar Pradesh, west India. This custom of buckwheat use as a fresh vegetable is found both in eastern India and western India. 4-2. Common buckwheat in India and in the Hima- layan hills is a short day-length plant Common buckwheat in India and in the Himalayan hills is usually cultivated in fall to early winter, from Sep- tember to December. As a result of cultivation under the condition of mild temperature and of short day-length, buckwheat in India and the Himalayan hills have become short day-length plant, with characteristics of tall vigor- ous vegetable parts with relatively longer cultivation pe- riod as compared with buckwheat from northern China and Japan. European common buckwheat has the characteristics of long-day to neutral day-length plant as the descend- ants of diffused buckwheat through the Silk Road (Ohni- shi, 1993, 2016). The characteristics of buckwheat in the Himalayan hills, short-day plant, is opposite to the char- acteristics of European buckwheat, day-neutral to long day-length plant. This leads to the conclusion that Euro- pean buckwheat does not have the origin in India nor in the Himalayan hills. Figure 3. Diffusion rout of culti- vated buckwheat from Mankang district in the Sanjiang area to India and the Himalayan hills. 61 Fagopyrum 41 (2): 57-63 (2024) 4-3. Making buckwheat noodle by hands in Ladakh of India Buckwheat noodle was not developed well in Nepal, India, and Pakistan. Two methods of making buckwheat noodle (Ohnishi, 2016), one using a noodle making wooden equipment, I call this as a Chinese method, one using special cooking knife to cut and make fine noodle of buckwheat dough. I call this as a Japanese method. Neither methods diffused to Nepal and India. In Bhu- tan a buckwheat noodle making equipment, called Put- ta in Bhutan, is used, hence buckwheat noodle is served as a dairy food. If peoples know neither methods, what happens for them? In a section of this symposium, Mr. Inazawa and his group will report the method for making buckwheat noodle by hands in Ladakh of India, where peoples make buckwheat noodle by their own hands only without using any special equipments, such as putta in Bhutan or special kitchen knife as in Japan. 5. REFERENCES Cheng, Q. F., 1999. 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Nei, 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Mol. Biol. Evol. 4: 405-425. 62 Ohnishi (2024): The Origin of Cultivated Buckwheat Swofford, D. L., 1999. PAUP*: Phylogenetic analysis using parsimony (* and other methods). Version 4.0b4a. Sinauer, Sunderland, Mass. USA. Swofford, D.L., 2002. PAUP*. Phylogenetic analysis using parsimony (* and other methods). Version 4.0. Sinauer, Sun- derland, Mass. USA. Wang, T. 1986. Genetic resources in Tibet. Variety Resources of Crops 1986(2), 23-25. (in Chinese) IZVLEČEK Izvor gojene ajde na območju distrikta Mankang območja Sanjiang Vzhodnega Tibeta in razširitev v Indijo ter na območje Himalaje Iskali in zbirali so naravne populacije divjega prednika gojene navadne ajde, začeli so leta 1990 in zaključili 2005. Med zbranimi vzorci je bil vzorec Zhuka, Xihe iz Mankanga, Tibet, najbolj soroden gojeni navadni ajdi. Po drugi strani, vzorca gojenih ajd iz Zhouba in Zhubalonga, oboje iz Mankanga, sta najbolj sorodna divjemu predniku navadne ajde. Na osnovi tega lahko oblikujemo hipotezo o izvoru gojene ajde v Mankangu na območju Sanjianga. Pot širjenja od prvot- nega izvora v Indijo in na območja gorovja Himalaje je predlagana v tej razpravi. Avtor opisuje lastnosti navadne ajde v Indiji in na območju Himalaje. Glavni zaključek je, da ajda v Evropi ne izvira iz Indije ali z območja Himalaje. Verjetno je v Evropo prišla iz Severne Kitajske po Svilni poti. 63 Fagopyrum 41 (2): 57-63 (2024)