Ethnobotanical review of traditional use of wild food plants in Japan

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However, there are fewer studies on the use of wild food plants in Japan than in other parts of the world. This research aims to provide an ethnobotanical review of the traditional food use of wild plants in Japan. It draws upon the largest database of traditional Japanese foodways, with a specific focus on the period approximately 1930. This occurred prior to Japan’s rapid economic growth, which resulted in significant changes in the use of wild plants. Methods The use of wild food plants in Japan was reviewed by studying the collection of Japanese foodways ( Nihon no shokuseikatsu zenshu ). The review extracted the local and common names, use locations, uses, processing and cooking methods, habitats and other relevant details pertaining to the reported use of wild food plants. This information was subsequently analyzed via Excel. Results A total of 268 taxa belonging to 87 families used in Japan approximately 1930 were documented. Asteraceae was the most frequently reported family as well as the most species-rich family. In terms of use, vegetables composed the largest group, followed by dessert fruits. The majority of species have been documented in only a limited number of reports. The review revealed local traditional uses of plants, including the use of plants as binders for making rice cake, the selection of plants according to their sex, and techniques for detoxifying toxic plants. Conclusion A comparison with those of previous studies on wild species used in other Asian regions revealed a similar preference for vegetables. However, notably, in Japan, ferns are also consumed in considerable quantities. Conversely, further elucidation is clearly needed with respect to certain matters, including the perception of plant sex and the relationships between diverse ash components and detoxification techniques. It is evident that further ethnobotanical research is needed, both in the form of an examination of ethnographic records and in the field, to gain a deeper understanding of the use of wild food plants in Japan. Wild food plants food wild species traditional knowledge Japan Asia Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Among more than 50 000 edible plant species in the world, only nine account for more than 66% of all crop production ( 1 ). On the other hand, there are diverse land- and culture-based food uses of wild plants in different parts of the world. Understanding the patterns of use and cultural significance and value of wild food plants worldwide has been important in terms of ecology, culture, nutrition, economics and agriculture, as well as in terms of genetic resources and biocultural diversity( 2 – 5 ). Previous ethnobiological studies have investigated the use of wild food plants in different populations around the world. In Europe in particular, there is an accumulation of ethnobotanical studies and reviews in many regions and countries( 6 – 14 ). With respect to Asia, there are studies on plant use and traditional knowledge, mainly in China, Southeast Asia and Central Asia, covering various social populations, such as farmers, indigenous communities and ethnic minorities( 15 – 26 ). Japan, the subject of this research, is located off the northeast seaboard of the Eurasian continent and has a surface area of nearly 378000 km2. The climate in Japan is diverse, with variations from subarctic in the northern regions to subtropical in the southern areas. A total of 8118 species of wild vascular plants are known to be grown in Japan( 27 ), with more than 1,000 of these species considered edible( 28 ). The use of wild food plants and mushrooms has been the subject of considerable interest in various academic disciplines in Japan. This has focused on a range of topics, including natural resources, nontimber forest products (NTFPs) and forest management in forestry( 29 – 31 ); the distribution of natural resources and territory in human geography( 32 ); and the practice of gathering and understanding human–nature relations in folklore studies and anthropology( 33 – 36 ). Their uses in Japan, as in other global contexts, were driven primarily by subsistence needs. Nevertheless, they also serve as intermediaries for communication and social interaction, particularly in the form of gift-giving practices, and some also serve as special food for ceremonies and special occasions, including the new year’s celebration and the Bon festival ( 31 , 37 ). Several previous studies in Japan have employed a variety of methods, including questionnaires, field surveys, and literature reviews, to ascertain the actual pattern of the use of wild food plans in each prefecture and region. For example, a study conducted in Nagano Prefecture revealed the use of 161 species of wild vegetables and 100 species of wild fruits in traditional foods between 1868 and 1955( 38 ). A more recent study conducted in the Noto Peninsula, Ishikawa Prefecture, revealed the presence of 176 species belonging to 65 families of edible plants( 39 ). However, the number of species of wild plants actually consumed was limited. Another study conducted in a village within the same Noto Peninsula identified the use of 21 species of wild food plants( 40 ). In Fukushima Prefecture in the Tohoku Region, a total of 45 taxa of wild vegetables and 26 taxa of nuts were used; however, these species have been used less frequently in recent years( 41 ). On the other hand, research on the use of wild food plants at the national scale is limited; however, a questionnaire survey conducted approximately 1980 revealed that more than 250 species of wild vegetables were used in Japan, with the exception of Okinawa ( 42 ). Another study suggested that 178 species from 46 families of vascular plants and 10 species of ferns are consumed as wild plants in northern Japan( 35 ). However, these include those that grow wild only in limited areas, such as Hokkaido, coastal regions and high-altitude areas, and those that grow wild throughout the country but are eaten only in certain regions. The same study also revealed that 101 species of seeds, including nuts and fruits belonging to 34 families, are consumed for medicinal and food purposes. It is suggested that the use of wild food plants is declining in Japan as a whole( 31 ). In particular, the postwar period of rapid economic growth from the late 1950s onward saw a shift in the use of wild food plants( 31 , 34 , 41 , 43 ). During this period, the social and natural environment, including industrial patterns, lifestyles and land use, underwent substantial transformation( 44 ). The study in Fukushima considered several specific factors that have contributed to the decline in the use of wild food plants( 41 ). These include social changes, such as changing lifestyles and the introduction of purchasable foods, and environmental changes, including land development, the use of pesticides, and the reduction in habitats. Conversely, since the period of economic growth, the popularity of gathering and consuming wild food plants has increased ( 45 ). This popularity is referred to as the sansai (mountain vegetable) boom and can be considered the period when the new term ‘ sansai ’ to describe mountain vegetables spread, especially in urban areas. There are also reports of new species being used as a result of access to new information( 41 ) and recreational foraging as a new trend( 31 ). An international review noted that most studies on gatherings are from the Americas, Europe, Central Asia and Africa, and there are fewer literatures and studies available from Asia( 3 ). Furthermore, most of the previous studies on the Japanese use of wild food plants were published in Japan and therefore limited accessibility to an international audience. Additionally, the majority of these studies were based on research conducted in specific geographical regions, such as prefectures or villages. To address this gap, this research aims to provide an ethnobotanical review of the traditional food use of wild plants in Japan via the largest database of traditional Japanese foodways, with a specific focus on approximately 1930, which was before rapid economic growth, when major changes in the use of wild plants occurred. The specific focuses of this study were ( 1 ) to identify how many species of plants were used as food in Japan around 1930, ( 2 ) to analyze which plant families and species were commonly used in local food categories, and ( 3 ) to understand the cultural uses of wild species in the context of Japanese traditional food culture. Methods Materials The use of wild food plants in Japan was reviewed by studying the collection of Japanese foodways ( Nihon no shokuseikatsu zenshu )( 46 ). The collection aimed to archive Japanese traditional foodways on the basis of an oral history of dietary habits during the late 1920s and early 1930s. This is the largest database on Japanese traditional foodways and contains records of approximately 52,000 dishes, including daily recipes as well as culinary uses of wild plants. The data were collected from interviews with 5,000 speakers who were mostly women in their late 70 s or older at the time of the survey and were involved in food preparation around the late 1920s to early 1930s. The collections consisted of 50 volumes covering all 47 prefectures from Hokkaido to Okinawa as well as one indigenous population, Ainu, in Hokkaido. This review examined 47 volumes, one for each prefecture (Fig. 1). The review extracted the local and common names, use locations, uses, processing and cooking methods, habitats and other relevant details pertaining to the reported use of wild food plants. This information was subsequently analyzed via Excel. Figure 1. Locations of regions and prefectures in Japan Wild species In this study, wild plants refer to noncultivated vascular plants, as suggested by a previous study( 7 ). Only species that could be identified as noncultivated plants were extracted on the basis of their description, including their habitats, such as mountains, fields, and river margins, as well as their growth conditions. In certain instances, both wild and cultivated forms, such as Eutrema japonicum , Diospyros kaki , Castanea crenata , and Phyllostachys edulis , can be found growing in the same study area. Only those species that are explicitly described as wild forms were included. Species that grow in areas with some degree of human intervention, such as the banks of rice paddies, or those that were cultivated but have escaped, such as Eleutherococcus sieboldianus , are also included if they are described as wild plants in the literature. In the collections, the local and Japanese common names of the plants are provided, with the exception of those for which only one name is available. In cases where only the local name was known, the common name was identified by consulting the Collection of Plants Dialects of Japan( 47 ) and the Folklore Research Card Collection Database, which was collected by Tama Saito, an independent folklore researcher who collected folklores that included uses of plants throughout Japan( 48 ), and other literature specific to each study area ( 49 – 54 ). A checklist of Japanese plant names( 55 ) was used to check the distribution of the species as well as to determine the scientific name from the Japanese name. The scientific name follows Plants of the World Online( 56 ), and synonyms have been corrected. Categories First, the present study included plants that are consumed as food or beverages, while plants used only as famine food or medicines were excluded. In accordance with the Economic Botany Data Collection Standard( 57 ), the uses of plants were first categorized into the following food categories: cereals, pseudocereals, pulses, nuts, dessert fruits, vegetables (including green and root/tuber vegetables), leaf protein concentrates, starches, oils/fats, gums/mucilages, sugars, and other food types. The vegetable category also included plants that are used in relatively small quantities and serve as flavor-adding seasonings. Among these categories, the categories of cereals, pseudocereals, pulses, leaf protein concentrates, gums/mucilages, and sugars were excluded, as there were no reports in these categories. In addition, by referring to folk perceptions of food uses, snacks, beverages, and binders were added as categories. Plants eaten between or after meals, with the exception of dessert fruits, were classified as snacks. Beverages include both alcoholic beverages such as liquors and nonalcoholic beverages such as tea. Binders include plants that are used to combine food ingredients. This usage is called " tsunagi " in Japanese and is used to make local foods such as mochi (rice cakes), dango (rice dumplings), and soba (buckwheat noodles). In total, 9 types of food categories were established as follows: nuts, dessert fruits, vegetables, starches, oils/fats, snacks, beverages, binders, and other food types. The plant parts used were classified into the following categories: entire plants, unspecified aerial parts, galls, stems, bark, leaves, inflorescences, infructescences, seeds, roots, exudates, shoots, and propagules. When parts used were not specified in the literature, they were classified as N/A (not available). Data analysis In this study, a report refers to a citation of the use of a wild food plant species (or taxa) in the literature at a single study site. For example, at one study site, if the fruit of species A was eaten as a dessert fruit and the shoot was eaten as a vegetable, the number of reports was counted as one. On the other hand, when the analysis was conducted in relation to the food categories and parts used, the calculation was based on the number of species in each category. In other words, if species A was eaten as a vegetable or fruit, it was counted as one species in each food category. Some reports were able to be identified only at the genus or subfamily level. The reason for this was that a single folk name encompassed multiple species. For example, azami is a general term for Cirsium spp. (Asteraceae), similar to the English thistle, and there are more than 60–70 species of azami in Japan. Similarly, noichigo or kiichigo is a generic name for strawberry plants that grow wild. Most of them belong to Rubus spp. (Rosaceae), and there are more than 30 wild species of Rubus spp. in Japan. The diversity of folk taxa and their species are noteworthy( 6 ). However, to avoid confusion in calculations, these folk taxa identified to the genus or subfamily level were calculated as one taxon, only if no botanical species from the genus or subfamily were recorded at the study site. Results and Discussions General data Among the 310 study sites across Japan, 278 reported the food use of wild plants. The study sites where no use was reported were mostly urban and coastal areas. In total, 3699 reports were recorded, 36 of which could not be identified by scientific names. As a result, 251 species belonging to 86 families that were used as food and beverages approximately 1930 in Japan were recorded. In addition, some reports (456 reports) were able to be identified only at the genus or subfamily level. Among the 17 such folk taxa, 16 genera and 1 subfamily belonging to 13 families were identified. A total of 268 taxa belonging to 87 families were used in Japan, including those specified above. The wild food plants identified in this study, along with their Japanese common names, food types, parts used, common uses, and number of reports, are listed in the appendix. Overall, Asteraceae (643 reports) was the most frequently reported family, followed by Poaceae (320), Araliaceae (239), Apiaceae (227), and Dennstaeditiaceae (199). The most species-rich family was Asteraceae (29 species), followed by Rosaceae (28), Poaceae (17), Apiaceae (11), Ericaceae (10), and Fabaceae (10). Table 1 lists the species with more than 100 use reports in all Japanese regions. Among the 278 study sites, Petasites japonicus was used in 78%, Artemisia princeps in 75%, Pteridium latiusculum in 72% , Osmunda japonica in 58%, Oenanthe javanica in 50%, Castanea crenata in 44%, and Aralia cordata in 44%. All of these plants were eaten as vegetables except A. princeps , which was used as a binder. All seven species are still commonly consumed at present and are also found in both cultivated and wild forms. On the other hand, the majority of species were sparsely reported. Eighty-three species (31% of the total taxa) were reported only once, 38 (14%) were reported twice, and 33 (12%) were reported three times. Table 1. List of wild species traditionally used as food in Japan with more than 100 reports Species Family Common Name Number of Reports Petasites japonicus (Siebold & Zucc.) Maxim. Asteraceae fuki 218 Artemisia princeps Pamp. Asteraceae yomogi 209 Pteridium latiusculum (Desv.) Hieron. Dennstaedtiaceae warabi 199 Osmunda japonica Thunb. Osmundaceae zenmai 160 Oenanthe javanica DC. Apiaceae seri 138 Castanea crenata Siebold et Zucc. Fagaceae kuri 123 Aralia cordata Thunb. Araliaceae udo 121 Figure 2 shows choropleth maps of Japan showing the numbers of wild food plant reports (a) and species (b) in each prefecture. Gifu Prefecture in the Chubu region had the greatest number of species (71 species), followed by Okayama (62), Niigata (61), Hiroshima (57), and Shizuoka (55) (Fig. 2(a)). Concerning the number of reports, Gifu also had the highest number of reports (134 reports), followed by Niigata (123), Akita (120), Nagano (116), and Aomori (106) (Fig. 2(b)). Fig. 2 The number of reports and species of wild food plants used in each prefecture In previous studies, regional differences in the use of wild food plants in Japan have been discussed according to different forest zones(58, 59). Several studies have suggested that the gathering and consumption of wild food plants and mushrooms has been more active in deciduous broad-leaved forests (or beech ( Fagus crenata ) forest zones in Japanese terminology), which primarily cover eastern Japan, especially in the heavy snowfall area from the Tohoku to Chubu regions, than in evergreen broad-leaved forests (or lucidophyllous forest zones), which primarily cover western Japan(31, 33, 60, 61). As the number of study sites in each prefecture varies, the frequency and diversity of wild food plant use among prefectures are not comparable comprehensively with those in this study alone. Nevertheless, the findings of this study do not contradict those of previous studies. Food Category Among the 9 food categories that were considered, vegetables were the largest group with 146 species, followed by dessert fruits with 78, beverages with 23, and snacks with 19 (Fig. 3). Fig. 3 The percentage of species in each food category (n=268) The uses of most species (219 species, 82% of the total species) were limited to only one category, while 42 species were used for two categories, and 6 species were used for 3 categories. Only one species ( Vitis coignetiae) was used in four categories: binders, beverages, dessert fruits, and other food types. The young leaves of these plants are used as binders in the production of mochi cakes. The infructescences were eaten raw as dessert fruits and were also used to make various beverages, such as juice, homemade liqueurs, and wine. Young fruits and exudates from vines are used in the production of agar jelly called tokoroten, and fruits are used to make local fermented foods such as lactic fermented vegetables. Vegetables Wild vegetables are called sanssai (mountain vegetables) in Japan. However, the term sansai is believed to have been used since approximately 1960 (35). The folk nomenclature of wild vegetables varies across different regions. For example, they are referred to as aomono (green/blue things), yama no mono (mountain things), and so forth. Aomono , which can be directly translated as ‘blue things’, is a common name for vegetables and is used in the case of wild vegetables such as yama no aomono (green vegetables of mountains). The most represented families for vegetables were Asteraceae (24 species), followed by Poaceae (15), Apiaceae (11), and Brassicaceae (8) (Fig. 4 (a)). These plants were eaten mostly in various ways, including aemono (boiled and dressed with seasonings) and nimono (simmered dishes). Some vegetables consumed raw included those used in small quantities for flavoring and seasoning, such as Allium schoenoprasum and Zanthoxylum piperitum . Some species of Dioscorea , including D. polystachya and D. japonica , are also consumed raw in the form of grated roots, which are used in the preparation of tororo, a seasoned slimy dish. Pickling is also a common method of cooking and preservation, utilizing a variety of ingredients, including salt, miso, sake lees and rice bran. As the majority of the species were collected during the spring season, these pickles were consumed as preserved food and were valuable foodstuffs during the winter months when vegetables were scarce. Other methods of preservation have also been employed, such as sun drying, salting and simmering down with soy sauce and sugar, known as tsukudani . Some plants, such as Aster yomena and Circium spp., were incorporated into rice dishes, including porridge. These vegetables contribute to the overall texture and flavor while also increasing the volume of the dishes. The ingredients used to increase the volume of rice-based dishes are referred to as " kate " or “ katemono ”. Historically, these were regarded as famine foods(62). The use of species to increase the volume of rice dishes has been similarly reported, for example, the use of the roots of Dioscorea bulbifera in Thailand(19). Fig. 4 Botanical families with the greatest number of species used for the four major food categories. Percentages of botanical families for (a) vegetables (n=146), (b) dessert fruit (n=78), (c) beverages (n=23), and (d) snacks (n=19) The most common parts used as vegetables were shoots (61 species), which represent nearly half of the species used as vegetables. This was followed by leaves, unspecified aerial parts (33 species), roots (19) and stems (18). Five species of inflorescences were used as vegetables, with Petasites japonicus being the most commonly consumed throughout the country. The young flower buds are called fukinoto and are consumed in a variety of ways, including as tempura and cooked with miso. The fukinoto was described as a plant that sprouts first in spring and heralds the arrival of spring at several study sites. Other species whose inflorescences are eaten as vegetables include Wisteria spp. and Erythronium japonicum . Propagules of 3 species were used as vegetables during autumn. The propagules are called mukago in Japanese, and two species of mukago yams ( Dioscorea japonica and D. polystachya ) are commonly eaten. They were eaten by roasting, boiling or as the kate ingredients as mentioned previously. The propagules of Elatostema involucratum were also eaten, although this was limited to one report. The leaves of these plants were removed, and only the stems with propagules were marinated in miso paste. They continue to be consumed in modern times in the northern part of Japan, such as Akita Prefecture, and are popular due to their rarity. These results are consistent with those reported in previous studies, such as those in Thailand, Laos, and China, which demonstrated a preference for shoots and leaves(15-17, 19), also known as herbophilia(6). Additionally, ferns such as Pteridium latiusculum , Osmunda japonica , and Oncoclea struthiopteris are also commonly consumed as vegetables in Japan. On the other hand, prior research on the use of wild species in South Korea has indicated that cooked shoots of species from Asteraceae are more frequently consumed as vegetables during the spring season (63-65). Therefore, the findings of the present study were consistent with those previously reported from Korea in terms of the plant taxa and part used, the preparation status (i.e., cooked or uncooked), and the season of use (i.e., spring). Dessert Fruits In total, there were 78 species of wild plants eaten as dessert fruits, of which almost one-third of the dessert fruits belonged to the Rosaceae family, followed by Ericaceae (8), Elaeagnaceae (4), and Rhamnaceae (4) (Fig. 4(b)). The most popular wild fruit was Akebia spp. ( A. squinata and A. trifoliata ). Their arils were eaten raw sometime as children’s snacks. Other parts of Akebia are also consumed as vegetables and tea. The stems and buds were consumed as tea and vegetables, while the pericarps were eaten as vegetables in various ways, such as pickled and stir-fried. The second most popular wild fruit was Rubus spp. As mentioned in the Methods section, wild strawberries are called noichigo (wild strawberry) or kiichigo (tree strawberry), and there are more than 30 wild Rubus species in Japan, including R. crataegifolius , R. parvifolius and R. sieboldii . Their infructescences were eaten raw, in many cases as children’s snacks. However, boiled or pickled fruits are also eaten in some cases. For example, fruits of Pyrus pyrifolia are eaten mostly raw, but in some areas, they are boiled for consumption. There was also a record of fruits of Vaccinium oxycoccos being pickled in salt, then sprinkled with sugar and eaten with green tea in the mountainous Oze area in Gunma Prefecture. Beverages There were 23 species used as beverages, including both alcoholic and nonalcoholic drinks (Fig. 4 (c)). Alcoholic beverages were of two types: so-called wines, in which infructescences were naturally fermented, and so-called liqueurs, in which plants were soaked in spirits such as shochu . Nonalcoholic beverages included tea and juice. Sixteen species were used as teas, including five species of Fabaceae, namely, Vicia bungei and Wisteria spp. Most of them used their aerial parts, such as leaves, shoots and inflorescence, and were processed by steaming, drying or roasting. On the other hand, one study reported that roots were used as tea. The roots of Imperata cylindrica were sun-dried and drunk as tea, which was favored by elderly people because it improves the production of urine. Following the tea, seven species were used to make liqueces. Most of them use infructescences for liqueurs, except for the species Geranium thunbergii, whose aerial parts were used 1 . In contrast, only two species ( Vitis coignetiae and V. ficifolia ) were used for making wine. Although the berries were used, the method of making wine exhibited slight regional variations, including the collection of berries when they were ripe or after frost or fermentation with or without sugar. Snacks Snacks are plants eaten between meals, excluding dessert fruits and nuts. In total, 19 species were eaten as snacks, including 4 from Poaceae and three each from Ericaceae and Polygonaceae (Fig. 4 (d)). Many of these snacks were eaten raw as children's snacks. Many were aerial parts, such as shoots, leaves, stems, and flowers, and there were many records of children foraging freely for snacks in the fields and mountains. Among the snacks, many sour-tasting plants, such as Rumex japonicus and Japanese knotweed ( Reynoutria japonica ), were eaten raw. They were sometimes eaten with salt on the spot of foraging. Similarly, children eating these acidic-tasting species have been documented in various European countries, including Spain, Sweden, and Poland(8). There were other snacks that have a sweet taste, such as common grass ( Imperata cylindrica ), whose young ears and other parts were eaten; nectar from some flowers, such as Azalea; and the roots of Cinnamomum sieboldii . Another interesting finding was that leaf galls of Rhododendron spp. (Azalea gall) were eaten raw as snacks, although this finding was limited to three reports. Azalea leaf galls are caused by fungi, including Exobasidium spp. (66), and locally called mochi (rice cake), the disease is called mochi disease. There have been reports of the use of plant galls in other parts of the world, including as folk medicine and for dyeing(67-69). However, reports of Azalea galls as snacks for children appear to be limited to the present study. Starches The 17 species used as starches were largely divided into those that use roots and those that use seeds. The most common family associated with the roots was Liliaceae (4 species), whereas the most common family associated with the seeds was Fagaceae (4 species), which included 3 Quercus spp. ( Q. acuta , Q. gilva, and Q. glauca ). Various techniques were used to extract starches, such as pounding, grating, soaking in water and strain. These extracted starches were used for making rice cakes, noodles, porridges and jelly type foods. Two toxic plants ( Lycoris radiata and Cycas revolute ) were included in this category. Although it was limited to only one report, bulbs of L. radiata were used for making starches. L. radiata contains lycorine, which is commonly found in the Amaryllidaceae (70). People were aware of this toxicity; therefore, they took a long time to remove the poison by being exposed to water repeatedly over a long period of time. Another poisonous plant was Cycas revolute . Their seeds and stems were used for extracting starches, particularly from the Ryukyu and Amami Islands of southern Japan. Like the use of L. radiata , to remove toxins, including cycasin, the plant parts used were exposed to water repeatedly over time. In some areas, people ferment them, and some report the infestation of maggots as a sign of detoxification. Binders Plants used to combine food ingredients are categorized as binders or “ tsunagi ” in Japanese. They were mostly used for rice cakes ( mochi and dango ) and sometimes for soba (buckwheat noodles). Among the 13 species used in this category, there were 8 species of Asteraceae, including two Artemisia spp. ( A. princeps and indica ) and 3 Synurus spp. ( S. pungens, deltoides and excelsus ). Young leaves and shoots of these species were boiled and added to food, and their trichomes and fibers contribute to the binding of other ingredients. In addition, some binder plants, such as Artemisia spp. and Pseudognaphalium affine, also contribute to the aroma or color of the prepared food. Although it was limited to two reports, the bark and cortex of red pine ( Pinus densiflora ) were used to make rice cakes called matsukawa mochi (pine skin rice cakes). Compared with other parts of the world, A. spp . and other species have been used as ingredients for mochi in China and Korea (71-73). Some research has also indicated that the use of P. affine for mochi in Japan was originally introduced from China and was later replaced by the use of A. spp. as an alternative to P. affine (74). Nuts Among the 13 species consumed as nuts, Fagaceae presented the greatest number of species (6 species). Although most species were tree plants, the seeds of 3 species of aquatic plants were also eaten as nuts ( Trapa natans var. Bispinosa, Nelumbo nucifera, and Euryale ferox e ). These nuts were eaten raw, roasted, or boiled and sometimes as snacks. Oils/fats There were only 3 species used for oils/fats ( Terreya nucifera , Camellia japonica , and C. Sasanqua ). Oils were extracted from seeds of these species. Among the three species, C. japonica was the most frequently reported, although its use was limited to only 6 reports in western Japan, particularly in the Kyushu region. The other two species were reported only once each. These oils were considered valuable and were sometimes sold and given as gifts. Other Food Types This category included the １3 species used for coloring, flavoring, fermented foods, or algae jelly making. For coloring and flavoring, for example, Gardenia jasminoides was used to add yellow color to food, including rice cakes and pickled vegetables. Another example is chick weeds ( Stellaria media ). In addition to their use as vegetables, they were also used for coloring and flavoring. In the Nasu area of Tochigi Prefecture, their aerial parts were used along with shiso leaves ( Perilla frutescens ) to provide sourness and better color when pickling with plums ( Prunus mume ). There were 8 species used for fermented foods, and their reports were all regionally specific. Some species ( Humulus cordifolius and Equisetum arvense ) wereadded when alcoholic beverages such as rice wine were made since they were considered to accelerate fermentation in the Tohoku region. In Nagano Prefecture, juices made from infructescences of 3 species ( Malus toringo , Pyrus pyrifolia , and Cycas revoluta ) were used for unsalted lactic-fermented pickles called sunki . These compounds are called “sources” or “essences” of sunki pickles and are believed to contribute to pH control and the production of lactic acid, etc. (75). As mentioned previously, the use of Cycas revolute was limited to only southern islands. On Amami islands, seeds of C . revolute were also used to make koji molds (fermentation starters) for making fermented soy bean paste, miso . Two species ( Oxalis corniculata and Vitis coignetiae ) were used to make tokoroten , jelly noodles made from red algae, including Gelidium amansii . The use of O. corniculata was reported only once in Ishikawa Prefecture, while there were 3 reports of the use of V. coignatiae in the Tohoku and Hokkaido regions. The parts used and how to use them differ from each other. Aerial parts of O. corniculate or unripen fruits of V. coignetiae were used when boiling red algae instead of vinegar. These compounds are considered to help extract agarose from red algae(76). Sour-tasting saps from vines of V. coignetiae were also used after red algae were boiled, but they were used to curdle the agar jelly. Thus, these sour plants were used to extract or solidify the jelly. Toxic Plants There were several reports of the use of toxic plants, although the type and degree of toxicity vary. For example, species containing carcinogens were eaten. As mentioned, bracken ( Pteridium latiusculum ) is commonly eaten throughout Japan, with shoots being eaten as vegetables and bracken starch extracted from the roots. Bracken contains a carcinogen called ptaquiloside (77). Another plant that contains carcinogens is Cycas revolute. Stems and seeds of C. revolute were used to make starch and fermentation starters; however, C. revolute contains a carcinogen called cycasin, which is found in other cycads(78). Alkaloid-containing species were also consumed. For example, rhizomes of Dioscorea spp. (specifically D. tokoro ) were boiled and consumed. Their rhizomes contain dioscin, a type of saponin that is commonly found in Dioscorea spp. (79). It was initially reported to have a very bitter taste but became palatable once people became accustomed to it in Saga Prefecture. In two of the five reports of D. tokoro as food, they were also used as an offering. The use as offerings to the gods has been reported in several places in Japan(36). Another alkaloid-containing species consumed was Lycoris radiata for starch from its bulbs, as mentioned previously. One of the most toxic plants eaten is the lacquer tree ( Toxicodendron vernicifluum ), which widely grows in China, Korea and Japan. In Asia, sap is used as a lacquer, but it contains an allergen called urushiol(80). In Miyagi and Ibaraki prefectures, their shoots were consumed as vegetables. In Miyagi, older individuals liked to consume shoots because of their soft texture, whereas in Ibaraki, they were regarded as tasty but potentially irritating food, with the decision to consume them dependent on the individual. It has been reported that individuals engaged in lacquer production, such as artisans, adhere to the custom of eating lacquer plants as a means of reducing their susceptibility to lacquer rash in Japan(81). However, the two regions reported in this study were not lacquer-producing areas and therefore cannot be relevant to this case of lacquer eating for immunity. On the other hand, the consumption of leafy shoots of T. vernicifluum as vegetables has been reported among ethnic minority groups in China, such as Lemo and Dulong in Yunnan Province and Yi in Sichuan Province(22, 82, 83). To understand the food use of lacquers in Japan, it is necessary to consider the food use of lacquers in the wider context of East Asia. Processing Techniques The majority of plants, particularly vegetables, were consumed primarily in cooked form. However, a range of other processing techniques were also employed. In particular, almost 300 reports of akunuki, a food processing technique used to remove aku , which are unpalatable and undesirable substances or tastes such as harshness and astringency, existed. The akunuki methods include the use of various ingredients, such as ash, rice water (water used to wash rice), rice bran, baking soda, vinegar, and camellia leaves; prolonged soaking in water; prolonged burial in soil; fermentation; and a combination of these methods. Some of these techniques are capable of removing not only unpleasant tastes but also toxic substances from wild species. The most common akunuki method was the use of ashes (179 reports). In many cases, the plant parts were soaked in water with ash for a long time or boiled. Among the species that were pretreated with ashes, warabi bracken ( Pteridium latiusculum ) was reported most frequently (80 reports), followed by Artemisia princeps (31), Osmunda japonica (28) and Aesculus turbinate (22). These reports included detailed descriptions of the specific types of ash to be used, for example, ash from stoves and hearths, rice straw ash and wood ash. There were several reports of the use of specific types of ash, particularly in the pretreatment of Aesculus turbinate seeds when they weremixed with glutinous rice to produce rice cakes ( tochi mochi ). Among the 22 reports, more than half described specific types of ashes to be used, including those of hardwoods, mainly Quercus spp. The use of ash, which is alkaline, for plant detoxification is a well-known traditional method involving pH control (84). The reason for the use of hardwood ashes, such as those derived from Quercus spp., is likely due to the pronounced alkalinity of these ashes, which has a greater impact on detoxification(85). The second most common method used for akunuki was the use of rice bran (58 reports). The plant parts were boiled in water with rice bran or, alternatively, left over water after rising rice was used. This method is commonly employed for the preparation of a variety of bamboo shoots. The use of rice bran is likely to facilitate the adsorption of undesired substances(84). However, in other Asian countries where bamboo shoots are widely consumed, boiling in water or saltwater is the predominant pretreatment method (86). A previous study suggested that the method of using rice bran is a relatively recent phenomenon that emerged after the late Meiji period (1868–1912) (87). Medicinal plants Previous studies have demonstrated a blurred boundary between food and medicine in the use of wild plants(88, 89). This study also identified 28 plant species that were used for food, medicinal and health-promoting purposes (Table 2). These plants were consumed for the treatment of various health conditions, including detoxication, deworming, cough, fever, tinnitus, and dizziness. Some of them were consumed specifically by infants, children, pregnant women, postpartum women and elderly individuals. The most frequently occurring species were those belonging to the Asteraceae family (5 species), followed by those of Lamiaceae (3 species). The species that were consumed as medicines included those that were commonly eaten as food throughout the country. For example, there were three reports of juice extracted from buds or roots of Petasites japonicus being given to newborns to detoxify the embryotoxic ( taidoku )(90, 91) 2 . Another example was Osmunda japonica. In the Tohoku region, they are believed to be beneficial for consumption after childbirth, with the aim of improving blood circulation or cleansing blood. Conversely, in Niigata Prefecture, eating them after birth was prohibited because they believed that they could cause blood to become “rough”. There were also six species, including Artemisia princeps, that were consumed as herbal teas, which are generally regarded as beneficial for the body. Table 2. A list of species consumed as food as well as for medicinal and health-related purposes Scientific name Family Parts used and preparations Functions Actinidia polygama Actinidiaceae seeds stomach Tetragonia tetragonoides Aizoaceae leaves gastrointestinal Oenanthe javanica Apiaceae shoots, boiled and mixed with miso or soy-sauce detoxication Artemisia indica Asteraceae juice from aerial parts fever Artemisia princeps Asteraceae herbal tea n/a Aster yomena Asteraceae shoots, boiled and mixed with miso or soy-sauce detoxication Circium spp. Asteraceae leaves, shoots n/a Crepidiastrum lanceolatum Asteraceae brew herbal tea fever Farfugium japonicum Asteraceae stems, boiled or salted detoxication Petasites japonicus Asteraceae juice roots detoxication particularly for newborn(roots) Pseudognaphalium affine Asteraceae brew herbal tea cough Platycodon grandiflorus Campanulaceae roots, pickled cough Elaeagnus umbellata Elaeagnaceae herbal tea tinnitus, deafness Pueraria montana var. lobata Fabaceae drink hot water mixed with starch extracted from roots food for the sick Geranium thunbergii Geraniaceae n/a n/a Clerodendrum trichotomum Lamiaceae shoots, cooked deworming, prevent summer fatigue Erythronium japonicum Liliaceae drink hot water mixed with starch extracted from roots food for the sick Osmunda japonica Osmundaceae shoots, cooked cleansings of blood during childbirth Plantago asiatica var. asiatica Plantaginaceae shoots, boiled and mixed with miso or soy-sauce, herbal tea detoxication Imperata cylindrica Poaceae roots, brewed as herbal tea promoting urination Hovenia dulcis Rhamnaceae infructescences, dried n/a Pyrus pyrifolia Rosaceae riped infructescences, raw cough, prevent asthma Phellodendron amurense Rutaceae debarked stems, brewed stomachach Houttuynia cordata Saururaceae herbal tea n/a Alkekengi officinarum Solanaceae infructescences, raw n/a Torreya nucifera Taxaceae seeds deworming Camellia japonica Theaceae oils from seeds deworming for children Vitis coignetiae Vitaceae infructescences, wine dizziness, good for health in general and for pregnant woman Male and Female Plants There were 5 reports on the uses of male and female plants of the 4 species. In Gifu Prefecture, stems of Reynoutria japonica were eaten raw as snacks with salt. The stems of the female plants were preferred because they were thicker, softer, and tasted sour, whereas the stems of the male plants were thinner and harder. Additionally, some reports indicate that plant sex determines whether a plant is used. For example, in Amami Ōshima Island of Kagoshima Prefecture, stems of Cycas revol uta were processed and used as starches. Only male plants were selected for starch extraction, as male plants were considered to store more starch. Another example involves approximately 2 species of Liliaceae ( Cardiocrinum cordatum and Lilium auratum var. auratum ), where the roots of the female plants were only selected for starch extraction. Reynoutria japonica and Cycas revoluta are dioecious plant species, whereas Lilium auratum var. auratum and Cardiocrinum cordatum are not. Thus, preferences for texture and flavor and whether to use plants were determined according to the sex of the plants. However, how male and female plants were locally identified for these 4 species was not described in the literature. As there are only a few studies on traditional knowledge of plant dioecy and the folk concepts of plant sex, further research is needed on the local perceptions of plant sex, as suggested by previous studies(92, 93). Taste and flavor There were a number of descriptions pertaining to the taste and flavor of wild species. These can be broadly classified into those pertaining to flavor, texture, and taste. Flavors were described as being of good quality, specific to wild species, in the spring, with all of them being perceived as positive. For example, in Akita, Hasteola robusta was carefully cooked as tempura to retain its distinctive aroma, which is considered important. There were several reports about texture, including the sliminess of Elatostema involucratum , the crispness of Astilbe rubra , and the stickiness and gooeyness of L. auratum var. auratum . These attributes were all regarded as positive. In terms of taste, wild species were often described as having a distinctive and slightly bitter taste, which wasperceived as good. This was exemplified in the preparation of seeds of Aesculus turbinata and bulbs of Lilium auratum var. auratum , where some described taking care not to lose all of the bitterness. There were also nine reports that the taste of wild species was better than that of cultivated or purchased species or those grown in proximity to their residences. Examples of these include udo ( Aralia cordata ), butterbur ( Petasites japonicus ), horse chestnut ( Castanea crenata ) and yam ( Dioscorea japonica ). Several previous studies have shown that wild vegetables were appreciated and preferred because of their tastes, for example, in southern Italy, Thailand, Himalaya and Japan (39, 94-97). Wild food plants were considered not only a source of nutrition or a means of sustenance but also a source of enjoyment through their unique tastes and flavors in Japan. Use of rare species There were few reports on the use of threatened species according to their current conservation status. Among the 2163 species of vascular plants listed on the 2020 Japanese Red List(98), the following four species were used approximately 1930: Platycodon grandifloras , prickly waterlily ( Euryale ferox ), and bird’s nest fern ( Asplenium antiquum ), which are listed as endangered, and Cinnamomum sieboldii , which is listed as near threatened. The roots of P. grandifloras were pickled in plum vinegar and were traditionally consumed in Hiroshima Prefecture. They are also used as a cough medicine, although their taste was not very tasty. There were 2 reports about the use of Euryale ferox . The cooked stems were eaten as vegetables, whereas the roasted or boiled seeds were eaten as snacks. Additionally, dried and ground seeds were used as starches to make dumplings ( dango ). Asplenium antiquum was reported once in Ishigaki city in Okinawa, where the shoots were used as vegetables in simmered or dressed dishes. Cinnamomum sieboldii was reported twice. The roots were eaten raw as snacks for the children. Conclusions This study examined Japan's largest food culture database and reviewed the food use of wild plants in the early Showa period (1926–1989). At least 268 taxa of plants belonging to 88 families were used to make food and beverages in Japan. Asteraceae was the most frequently reported and most species-rich family. The majority of species was documented in only a limited number of reports, and the number of species used at the national level was limited. In other words, each region and locality had its own distinctive culinary traditions involving wild food species. A comparison with those of previous studies on wild species used in other Asian regions revealed a similar preference for vegetables, namely, shoots and leaves. However, notably, in Japan, ferns were also consumed in considerable quantities. Furthermore, there were some other similarities with other Asian regions, including folk uses of plants such as binder plants and kate plants and the consumption of toxic plants (lacquers). Conversely, further elucidation is clearly needed with respect to certain matters, including the perception of plant sex, the relationship between the consumption of lacquer and immunity, and the relationships between diverse ash components and detoxification techniques. It is therefore evident that further ethnobotanical research is needed, both in the form of an examination of ethnographic records and in the field, to gain a deeper understanding of the use of wild food plants in Japan. Declarations Ethics approval and consent to participate Not Applicable Consent for publication Not Applicable Competing interests The authors declare that they have no competing interests Funding Lotte Foundation Author Contribution The author confirms sole responsibility from conceptualizing and designing study , data collection, analysis and interpretation of results, and to manuscript preparation. Acknowledgement This research was supported by JSPS KAKENHI Grant Numbers JP21H03685, JP21K01086, and Lotte Foundation. Availability of data and materials I confirm that the data supporting the findings of this study are available within the article and its supplementary material. References Bélanger J, Pilling D. 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Bulletin of Kagoshima Women's Junior College. 2004;39:21-5. Chongtham N, Bisht MS, Premlata T, Bajwa HK, Sharma V, Santosh O. Quality improvement of bamboo shoots by removal of antinutrients using different processing techniques: A review. Journal of Food Science and Technology. 2022:1-11. Iwamura M, Miyaka N, editors. Takenoko no akunuki ni kansuru kenkyu (Research on akunuki of bamboo shoots) (in Japanese). The Japan Society of Cookery Science; 2015; Shizuoka, Japan. Pieroni A, Price L. Eating and healing: traditional food as medicine: Crc Press; 2006. Etkin NL. Medicinal cuisines: diet and ethopharmacology. International Journal of Pharmacognosy. 1996;34(5):313-26. Daidoji K. The adaptation of the treatise on cold damage in eighteenth-century Japan: Text, society, and readers. Asian Medicine. 2013;8(2):361-93. Triplett K. Pediatric Care and Buddhism in Premodern Japan: A Case of Applied “Demonology”? Asian Medicine. 2020;14(2):313-41. Seethapathy GS, Ravikumar K, Paulsen BS, de Boer HJ, Wangensteen H. Ethnobotany of dioecious species: Traditional knowledge on dioecious plants in India. J Ethnopharmacol. 2018;221:56-64. Berlin B, Breedlove DE, Raven PH. General Principles of Classification and Nomenclature in Folk Biology. American Anthropologist. 1973;75(1):214-42. Ghirardini MP, Carli M, del Vecchio N, Rovati A, Cova O, Valigi F, et al. The importance of a taste. A comparative study on wild food plant consumption in twenty-one local communities in Italy. J Ethnobiol Ethnomed. 2007;3:22. Somnasang P, Moreno-Black G. Knowing, gathering and eating: knowledge and attitudes about wild food in an Isan village in Northeastern Thailand. Journal of Ethnobiology. 2000;20(2):197-216. Thakur D, Sharma A, Uniyal SK. Why they eat, what they eat: patterns of wild edible plants consumption in a tribal area of Western Himalaya. Journal of ethnobiology and ethnomedicine. 2017;13:1-12. Sugiura T, Kishimoto K. On the liking of wild vegetables and mushrooms by consumers. Journal of the Japanese Forestry Society. 1989;71(1):31-8. Environment JMot. Japanese Red List 2020. In: Environment JMot, editor. 2020. Footnotes G. thunbergii was a common folk medicine in Japan. A number of reports in the collection also showed that they were used as a medicine. As explained in the Methods, plants that were only used as medicines were not included in this study. This particular report was included since G. thunbergii was described as a wild plant used to make herbal liquors which had both food and medicinal purposes. At the time, it was believed that taidoku could cause neonatal illnesses such as boil and rash, and plants and medicines were given to infants shortly after birth. This theory of embryonic poisoning is thought to have originate in China. Additional Declarations No competing interests reported. Supplementary Files Appendix.xlsx Cite Share Download PDF Status: Published Journal Publication published 21 Nov, 2024 Read the published version in Journal of Ethnobiology and Ethnomedicine → Version 1 posted Editorial decision: Revision requested 10 Aug, 2024 Reviews received at journal 10 Aug, 2024 Reviews received at journal 07 Aug, 2024 Reviewers agreed at journal 31 Jul, 2024 Reviewers agreed at journal 31 Jul, 2024 Reviewers invited by journal 31 Jul, 2024 Editor assigned by journal 30 Jul, 2024 Submission checks completed at journal 30 Jul, 2024 First submitted to journal 25 Jul, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4803162","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":338589819,"identity":"0a5a7744-fcb2-47b6-ac41-de98d307b584","order_by":0,"name":"Yoshimi Osawa","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA/klEQVRIiWNgGAWjYNCCCnQBCYJazpCshbGNaPcAAT//4WcSP+fV2vM3MD98zFOzTZ6B/fADBssduLVIzkgzk+zddjxxxgE2Y2OeY7cNG3jSDBgkMZyKAAY3GMxu8G47lmDAwGAmzcN2m7GBIQdoEh6nGpw//u3m3znH7A0Y2L9J8/y7bd/A/4aAlgM5Zrd5G2oYNzDwmEnztt1ObJAgYIvkjJzy3zLHDiTOOMxTbDi373Zym8QzgwP4/MLPf3yz4ZuaOnv+9vaND958u23bz5/88LEknhCDgsMMDMwMDEw8QCYbiCvZQFBLHZhk/AHlMn4krGUUjIJRMApGDgAA4AZPSKWkKk4AAAAASUVORK5CYII=","orcid":"","institution":"Aoyama Gakuin University","correspondingAuthor":true,"prefix":"","firstName":"Yoshimi","middleName":"","lastName":"Osawa","suffix":""}],"badges":[],"createdAt":"2024-07-25 16:06:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4803162/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4803162/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1186/s13002-024-00736-2","type":"published","date":"2024-11-21T15:57:06+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":63359162,"identity":"081f6f3f-f48c-4b92-9d6d-fdfccc8e7ae1","added_by":"auto","created_at":"2024-08-27 09:53:19","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":150436,"visible":true,"origin":"","legend":"\u003cp\u003eLocations of regions and prefectures in Japan\u003c/p\u003e","description":"","filename":"FIG1.png","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/6a4b9fbe874bec658b2f836f.png"},{"id":63358549,"identity":"20b620ca-50ae-4cff-a2b6-391608b565e3","added_by":"auto","created_at":"2024-08-27 09:45:19","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":24220,"visible":true,"origin":"","legend":"\u003cp\u003eThe number of reports and species of wild food plants used in each prefecture\u003c/p\u003e","description":"","filename":"FIG2.png","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/eb28afa5cf8d56a94adfb971.png"},{"id":63358550,"identity":"739e99f0-9f7f-40c4-8882-442a269cd541","added_by":"auto","created_at":"2024-08-27 09:45:19","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":13055,"visible":true,"origin":"","legend":"\u003cp\u003eThe percentage of species in each food category (n=268)\u003c/p\u003e","description":"","filename":"FIG3.png","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/b4f2ade289d7e8d68fa222bf.png"},{"id":63358551,"identity":"8ebceb4f-8354-49fd-a7fd-cc0b1b834dd1","added_by":"auto","created_at":"2024-08-27 09:45:19","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":37122,"visible":true,"origin":"","legend":"\u003cp\u003eBotanical families with the greatest number of species used for the four major food categories. Percentages of botanical families for (a) vegetables (n=146), (b) dessert fruit (n=78), (c) beverages (n=23), and (d) snacks (n=19)\u003c/p\u003e","description":"","filename":"FIG4.png","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/3533010a2d9a8f06c919bacc.png"},{"id":69835015,"identity":"f84f555c-6cfd-4d43-b7d1-b9a7226bad3c","added_by":"auto","created_at":"2024-11-25 16:11:36","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":958781,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/aa3767f3-fe75-45b5-bed3-d28eb0266544.pdf"},{"id":63358552,"identity":"0ef7a3bb-3e1b-4fcc-bad3-1adfbb9f4a4b","added_by":"auto","created_at":"2024-08-27 09:45:19","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":36215,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-4803162/v1/2e2b0e1613225539d4e589d4.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Ethnobotanical review of traditional use of wild food plants in Japan","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAmong more than 50 000 edible plant species in the world, only nine account for more than 66% of all crop production (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e). On the other hand, there are diverse land- and culture-based food uses of wild plants in different parts of the world. Understanding the patterns of use and cultural significance and value of wild food plants worldwide has been important in terms of ecology, culture, nutrition, economics and agriculture, as well as in terms of genetic resources and biocultural diversity(\u003cspan additionalcitationids=\"CR3 CR4\" citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e). Previous ethnobiological studies have investigated the use of wild food plants in different populations around the world. In Europe in particular, there is an accumulation of ethnobotanical studies and reviews in many regions and countries(\u003cspan additionalcitationids=\"CR7 CR8 CR9 CR10 CR11 CR12 CR13\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e). With respect to Asia, there are studies on plant use and traditional knowledge, mainly in China, Southeast Asia and Central Asia, covering various social populations, such as farmers, indigenous communities and ethnic minorities(\u003cspan additionalcitationids=\"CR16 CR17 CR18 CR19 CR20 CR21 CR22 CR23 CR24 CR25\" citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eJapan, the subject of this research, is located off the northeast seaboard of the Eurasian continent and has a surface area of nearly 378000 km2. The climate in Japan is diverse, with variations from subarctic in the northern regions to subtropical in the southern areas. A total of 8118 species of wild vascular plants are known to be grown in Japan(\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e), with more than 1,000 of these species considered edible(\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe use of wild food plants and mushrooms has been the subject of considerable interest in various academic disciplines in Japan. This has focused on a range of topics, including natural resources, nontimber forest products (NTFPs) and forest management in forestry(\u003cspan additionalcitationids=\"CR30\" citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e); the distribution of natural resources and territory in human geography(\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e); and the practice of gathering and understanding human\u0026ndash;nature relations in folklore studies and anthropology(\u003cspan additionalcitationids=\"CR34 CR35\" citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e). Their uses in Japan, as in other global contexts, were driven primarily by subsistence needs. Nevertheless, they also serve as intermediaries for communication and social interaction, particularly in the form of gift-giving practices, and some also serve as special food for ceremonies and special occasions, including the new year\u0026rsquo;s celebration and the Bon festival (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral previous studies in Japan have employed a variety of methods, including questionnaires, field surveys, and literature reviews, to ascertain the actual pattern of the use of wild food plans in each prefecture and region. For example, a study conducted in Nagano Prefecture revealed the use of 161 species of wild vegetables and 100 species of wild fruits in traditional foods between 1868 and 1955(\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e). A more recent study conducted in the Noto Peninsula, Ishikawa Prefecture, revealed the presence of 176 species belonging to 65 families of edible plants(\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e). However, the number of species of wild plants actually consumed was limited. Another study conducted in a village within the same Noto Peninsula identified the use of 21 species of wild food plants(\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e). In Fukushima Prefecture in the Tohoku Region, a total of 45 taxa of wild vegetables and 26 taxa of nuts were used; however, these species have been used less frequently in recent years(\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOn the other hand, research on the use of wild food plants at the national scale is limited; however, a questionnaire survey conducted approximately 1980 revealed that more than 250 species of wild vegetables were used in Japan, with the exception of Okinawa (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e). Another study suggested that 178 species from 46 families of vascular plants and 10 species of ferns are consumed as wild plants in northern Japan(\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e). However, these include those that grow wild only in limited areas, such as Hokkaido, coastal regions and high-altitude areas, and those that grow wild throughout the country but are eaten only in certain regions. The same study also revealed that 101 species of seeds, including nuts and fruits belonging to 34 families, are consumed for medicinal and food purposes.\u003c/p\u003e \u003cp\u003eIt is suggested that the use of wild food plants is declining in Japan as a whole(\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e). In particular, the postwar period of rapid economic growth from the late 1950s onward saw a shift in the use of wild food plants(\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e). During this period, the social and natural environment, including industrial patterns, lifestyles and land use, underwent substantial transformation(\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e). The study in Fukushima considered several specific factors that have contributed to the decline in the use of wild food plants(\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e). These include social changes, such as changing lifestyles and the introduction of purchasable foods, and environmental changes, including land development, the use of pesticides, and the reduction in habitats.\u003c/p\u003e \u003cp\u003eConversely, since the period of economic growth, the popularity of gathering and consuming wild food plants has increased (\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e). This popularity is referred to as the \u003cem\u003esansai\u003c/em\u003e (mountain vegetable) boom and can be considered the period when the new term \u0026lsquo;\u003cem\u003esansai\u003c/em\u003e\u0026rsquo; to describe mountain vegetables spread, especially in urban areas. There are also reports of new species being used as a result of access to new information(\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e) and recreational foraging as a new trend(\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAn international review noted that most studies on gatherings are from the Americas, Europe, Central Asia and Africa, and there are fewer literatures and studies available from Asia(\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e). Furthermore, most of the previous studies on the Japanese use of wild food plants were published in Japan and therefore limited accessibility to an international audience. Additionally, the majority of these studies were based on research conducted in specific geographical regions, such as prefectures or villages.\u003c/p\u003e \u003cp\u003eTo address this gap, this research aims to provide an ethnobotanical review of the traditional food use of wild plants in Japan via the largest database of traditional Japanese foodways, with a specific focus on approximately 1930, which was before rapid economic growth, when major changes in the use of wild plants occurred. The specific focuses of this study were (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) to identify how many species of plants were used as food in Japan around 1930, (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) to analyze which plant families and species were commonly used in local food categories, and (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) to understand the cultural uses of wild species in the context of Japanese traditional food culture.\u003c/p\u003e"},{"header":"Methods","content":"\u003ch2\u003eMaterials\u003c/h2\u003e\n\u003cp\u003eThe use of wild food plants in Japan was reviewed by studying the collection of Japanese foodways (\u003cem\u003eNihon no shokuseikatsu zenshu\u003c/em\u003e)(\u003cspan class=\"CitationRef\"\u003e46\u003c/span\u003e). The collection aimed to archive Japanese traditional foodways on the basis of an oral history of dietary habits during the late 1920s and early 1930s. This is the largest database on Japanese traditional foodways and contains records of approximately 52,000 dishes, including daily recipes as well as culinary uses of wild plants. The data were collected from interviews with 5,000 speakers who were mostly women in their late 70 s or older at the time of the survey and were involved in food preparation around the late 1920s to early 1930s. The collections consisted of 50 volumes covering all 47 prefectures from Hokkaido to Okinawa as well as one indigenous population, Ainu, in Hokkaido. This review examined 47 volumes, one for each prefecture (Fig.\u0026nbsp;1). The review extracted the local and common names, use locations, uses, processing and cooking methods, habitats and other relevant details pertaining to the reported use of wild food plants. This information was subsequently analyzed via Excel.\u003c/p\u003e\n\u003cp\u003eFigure\u0026nbsp;1. Locations of regions and prefectures in Japan\u003c/p\u003e\n\u003ch2\u003eWild species\u003c/h2\u003e\n\u003cp\u003eIn this study, wild plants refer to noncultivated vascular plants, as suggested by a previous study(\u003cspan class=\"CitationRef\"\u003e7\u003c/span\u003e). Only species that could be identified as noncultivated plants were extracted on the basis of their description, including their habitats, such as mountains, fields, and river margins, as well as their growth conditions. In certain instances, both wild and cultivated forms, such as \u003cem\u003eEutrema japonicum\u003c/em\u003e, \u003cem\u003eDiospyros kaki\u003c/em\u003e, \u003cem\u003eCastanea crenata\u003c/em\u003e, and \u003cem\u003ePhyllostachys edulis\u003c/em\u003e, can be found growing in the same study area. Only those species that are explicitly described as wild forms were included. Species that grow in areas with some degree of human intervention, such as the banks of rice paddies, or those that were cultivated but have escaped, such as \u003cem\u003eEleutherococcus sieboldianus\u003c/em\u003e, are also included if they are described as wild plants in the literature.\u003c/p\u003e\n\u003cp\u003eIn the collections, the local and Japanese common names of the plants are provided, with the exception of those for which only one name is available. In cases where only the local name was known, the common name was identified by consulting the Collection of Plants Dialects of Japan(\u003cspan class=\"CitationRef\"\u003e47\u003c/span\u003e) and the Folklore Research Card Collection Database, which was collected by Tama Saito, an independent folklore researcher who collected folklores that included uses of plants throughout Japan(\u003cspan class=\"CitationRef\"\u003e48\u003c/span\u003e), and other literature specific to each study area (\u003cspan class=\"CitationRef\"\u003e49\u003c/span\u003e\u0026ndash;\u003cspan class=\"CitationRef\"\u003e54\u003c/span\u003e). A checklist of Japanese plant names(\u003cspan class=\"CitationRef\"\u003e55\u003c/span\u003e) was used to check the distribution of the species as well as to determine the scientific name from the Japanese name. The scientific name follows Plants of the World Online(\u003cspan class=\"CitationRef\"\u003e56\u003c/span\u003e), and synonyms have been corrected.\u003c/p\u003e\n\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\n \u003ch2\u003eCategories\u003c/h2\u003e\n \u003cp\u003eFirst, the present study included plants that are consumed as food or beverages, while plants used only as famine food or medicines were excluded. In accordance with the Economic Botany Data Collection Standard(\u003cspan class=\"CitationRef\"\u003e57\u003c/span\u003e), the uses of plants were first categorized into the following food categories: cereals, pseudocereals, pulses, nuts, dessert fruits, vegetables (including green and root/tuber vegetables), leaf protein concentrates, starches, oils/fats, gums/mucilages, sugars, and other food types. The vegetable category also included plants that are used in relatively small quantities and serve as flavor-adding seasonings. Among these categories, the categories of cereals, pseudocereals, pulses, leaf protein concentrates, gums/mucilages, and sugars were excluded, as there were no reports in these categories.\u003c/p\u003e\n \u003cp\u003eIn addition, by referring to folk perceptions of food uses, snacks, beverages, and binders were added as categories. Plants eaten between or after meals, with the exception of dessert fruits, were classified as snacks. Beverages include both alcoholic beverages such as liquors and nonalcoholic beverages such as tea. Binders include plants that are used to combine food ingredients. This usage is called \u0026quot;\u003cem\u003etsunagi\u003c/em\u003e\u0026quot; in Japanese and is used to make local foods such as \u003cem\u003emochi\u003c/em\u003e (rice cakes), \u003cem\u003edango\u003c/em\u003e (rice dumplings), and \u003cem\u003esoba\u003c/em\u003e (buckwheat noodles). In total, 9 types of food categories were established as follows: nuts, dessert fruits, vegetables, starches, oils/fats, snacks, beverages, binders, and other food types.\u003c/p\u003e\n \u003cp\u003eThe plant parts used were classified into the following categories: entire plants, unspecified aerial parts, galls, stems, bark, leaves, inflorescences, infructescences, seeds, roots, exudates, shoots, and propagules. When parts used were not specified in the literature, they were classified as N/A (not available).\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\n \u003ch2\u003eData analysis\u003c/h2\u003e\n \u003cp\u003eIn this study, a report refers to a citation of the use of a wild food plant species (or taxa) in the literature at a single study site. For example, at one study site, if the fruit of species A was eaten as a dessert fruit and the shoot was eaten as a vegetable, the number of reports was counted as one. On the other hand, when the analysis was conducted in relation to the food categories and parts used, the calculation was based on the number of species in each category. In other words, if species A was eaten as a vegetable or fruit, it was counted as one species in each food category.\u003c/p\u003e\n \u003cp\u003eSome reports were able to be identified only at the genus or subfamily level. The reason for this was that a single folk name encompassed multiple species. For example, \u003cem\u003eazami\u003c/em\u003e is a general term for \u003cem\u003eCirsium\u003c/em\u003e spp. (Asteraceae), similar to the English thistle, and there are more than 60\u0026ndash;70 species of \u003cem\u003eazami\u003c/em\u003e in Japan. Similarly, \u003cem\u003enoichigo\u003c/em\u003e or \u003cem\u003ekiichigo\u003c/em\u003e is a generic name for strawberry plants that grow wild. Most of them belong to \u003cem\u003eRubus\u003c/em\u003e spp. (Rosaceae), and there are more than 30 wild species of \u003cem\u003eRubus\u003c/em\u003e spp. in Japan. The diversity of folk taxa and their species are noteworthy(\u003cspan class=\"CitationRef\"\u003e6\u003c/span\u003e). However, to avoid confusion in calculations, these folk taxa identified to the genus or subfamily level were calculated as one taxon, only if no botanical species from the genus or subfamily were recorded at the study site.\u003c/p\u003e\n\u003c/div\u003e"},{"header":"Results and Discussions","content":"\u003cp\u003e\u003cstrong\u003eGeneral data\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAmong\u0026nbsp;the 310 study sites across Japan, 278 reported\u0026nbsp;the\u0026nbsp;food use of wild plants.\u0026nbsp;The study\u0026nbsp;sites where no use was reported were mostly urban and coastal areas. In total, 3699 reports were recorded,\u0026nbsp;36\u0026nbsp;of which could not be identified by scientific names. As a result, 251 species belonging to 86 families that were used as food and beverages approximately 1930 in Japan were recorded. In addition, some reports (456 reports) were able to be identified only\u0026nbsp;at\u0026nbsp;the genus or subfamily level.\u0026nbsp;Among\u0026nbsp;the 17 such folk taxa, 16 genera and 1 subfamily belonging to 13 families were identified. A total of 268 taxa belonging to 87 families were used in Japan,\u0026nbsp;including those specified above. The wild food plants identified in this study, along with their Japanese common names, food types, parts used, common uses, and number of reports, are listed in the appendix.\u003c/p\u003e\n\u003cp\u003eOverall, Asteraceae (643 reports)\u0026nbsp;was the most frequently reported family, followed by Poaceae\u0026nbsp;(320), Araliaceae\u0026nbsp;(239), Apiaceae\u0026nbsp;(227), and Dennstaeditiaceae\u0026nbsp;(199).\u0026nbsp;The most species-rich family was Asteraceae (29 species), followed by Rosaceae (28), Poaceae (17), Apiaceae (11), Ericaceae (10), and Fabaceae (10).\u003c/p\u003e\n\u003cp\u003eTable 1 lists the species with more than 100 use reports in all Japanese regions. Among the 278 study sites, \u003cem\u003ePetasites japonicus\u003c/em\u003e was used in 78%, \u003cem\u003eArtemisia princeps\u003c/em\u003e in 75%, \u003cem\u003ePteridium latiusculum\u003c/em\u003e in 72%\u003cem\u003e,\u003c/em\u003e \u003cem\u003eOsmunda japonica\u003c/em\u003e in 58%, \u003cem\u003eOenanthe javanica\u003c/em\u003e in 50%, \u003cem\u003eCastanea crenata\u003c/em\u003e in 44%, and \u003cem\u003eAralia cordata\u003c/em\u003e in 44%. All of these plants were eaten as vegetables except \u003cem\u003eA. princeps\u003c/em\u003e\u003cem\u003e,\u003c/em\u003e which was used as a binder. All seven species are still commonly consumed at present and are also found in both cultivated and wild forms. On the other hand, the majority of species were sparsely reported. Eighty-three species (31% of the total taxa) were reported only once, 38 (14%) were reported twice, and 33 (12%) were reported three times.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 1. List of wild species traditionally used as food in Japan with more than 100 reports\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eSpecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eFamily\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eCommon Name\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of Reports\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003ePetasites japonicus\u003c/em\u003e (Siebold \u0026amp; Zucc.) Maxim.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003efuki\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e218\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eArtemisia princeps\u003c/em\u003e Pamp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eyomogi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e209\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003ePteridium latiusculum\u003c/em\u003e (Desv.) Hieron.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eDennstaedtiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003ewarabi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e199\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eOsmunda japonica\u003c/em\u003e Thunb.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eOsmundaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003ezenmai\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e160\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eOenanthe javanica\u003c/em\u003e DC.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eApiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eseri\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e138\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eCastanea crenata\u003c/em\u003e Siebold et Zucc.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eFagaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003ekuri\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e123\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eAralia cordata\u003c/em\u003e Thunb.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eAraliaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eudo\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003e121\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eFigure 2 shows choropleth maps of Japan showing the numbers of wild food plant reports (a) and species (b) in each prefecture. Gifu Prefecture in the Chubu region had the greatest number of species (71 species), followed by Okayama (62), Niigata (61), Hiroshima (57), and Shizuoka (55) (Fig. 2(a)). Concerning the number of reports, Gifu also had the highest number of reports (134 reports), followed by Niigata (123), Akita (120), Nagano (116), and Aomori (106) (Fig. 2(b)).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Fig. 2 The number of reports and species of wild food plants used in each prefecture\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn previous studies, regional differences in the use of wild food plants in Japan have been discussed according to different forest zones(58, 59). Several studies\u0026nbsp;have\u0026nbsp;suggested that the gathering and consumption of wild food plants and mushrooms has been more active in deciduous broad-leaved\u0026nbsp;forests\u0026nbsp;(or beech (\u003cem\u003eFagus crenata\u003c/em\u003e) forest zones in Japanese terminology), which primarily cover eastern Japan, especially in the heavy snowfall area from the Tohoku to Chubu regions, than in evergreen broad-leaved forests (or lucidophyllous forest zones), which primarily cover western Japan(31, 33, 60, 61). As the number of study sites in each prefecture varies, the frequency and diversity of wild food plant use among prefectures are not comparable comprehensively with those in this study alone. Nevertheless, the findings of this study do not contradict those of previous studies.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFood Category\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAmong the 9 food categories that were considered, vegetables were the largest group with 146 species, followed by dessert fruits with 78, beverages with 23, and snacks with 19 (Fig. 3).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFig. 3 The percentage of species in each food category (n=268)\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe uses of most species (219 species, 82% of the total species) were limited to only one category, while 42 species were used for two categories, and 6 species were used for 3 categories. Only one species (\u003cem\u003eVitis coignetiae)\u003c/em\u003e was used in four categories: binders, beverages, dessert fruits, and other food types. The young leaves of these plants are used as binders in the production of mochi cakes. The infructescences were eaten raw as dessert fruits and were also used to make various beverages, such as juice, homemade liqueurs, and wine. Young fruits and exudates from vines are used in the production of agar jelly called \u003cem\u003etokoroten,\u003c/em\u003e and fruits are used to make local fermented foods such as lactic fermented vegetables.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetables\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWild vegetables are called \u003cem\u003esanssai\u0026nbsp;\u003c/em\u003e(mountain vegetables) in Japan. However, the term \u003cem\u003esansai\u0026nbsp;\u003c/em\u003eis believed to have been used since approximately 1960 (35). The folk nomenclature of wild vegetables varies across different regions. For example, they are referred to as \u003cem\u003eaomono\u003c/em\u003e (green/blue things), \u003cem\u003eyama no mono\u003c/em\u003e (mountain things), and so forth. \u003cem\u003eAomono\u003c/em\u003e, which can be directly translated as \u0026lsquo;blue things\u0026rsquo;, is a common name for vegetables and is used in the case of wild vegetables such as \u003cem\u003eyama no aomono\u003c/em\u003e (green vegetables of mountains).\u003c/p\u003e\n\u003cp\u003eThe most represented families for vegetables were Asteraceae (24 species), followed by Poaceae (15), Apiaceae (11), and Brassicaceae (8) (Fig. 4 (a)). These plants were eaten mostly in various ways, including \u003cem\u003eaemono\u0026nbsp;\u003c/em\u003e(boiled and dressed with seasonings) and \u003cem\u003enimono\u0026nbsp;\u003c/em\u003e(simmered dishes). Some vegetables consumed raw included those used in small quantities for flavoring and seasoning, such as \u003cem\u003eAllium schoenoprasum\u003c/em\u003e and \u003cem\u003eZanthoxylum piperitum\u003c/em\u003e. Some species of \u003cem\u003eDioscorea\u003c/em\u003e, including \u003cem\u003eD. polystachya\u003c/em\u003e and \u003cem\u003eD. japonica\u003c/em\u003e, are also consumed raw in the form of grated roots, which are used in the preparation of \u003cem\u003etororo,\u003c/em\u003e a seasoned slimy dish. Pickling is also a common method of cooking and preservation, utilizing a variety of ingredients, including salt, miso, sake lees and rice bran. As the majority of the species were collected during the spring season, these pickles were consumed as preserved food and were valuable foodstuffs during the winter months when vegetables were scarce. Other methods of preservation have also been employed, such as sun drying, salting and simmering down with soy sauce and sugar, known as \u003cem\u003etsukudani\u003c/em\u003e. Some plants, such as \u003cem\u003eAster yomena\u003c/em\u003e and \u003cem\u003eCircium\u003c/em\u003e spp., were incorporated into rice dishes, including porridge. These vegetables contribute to the overall texture and flavor while also increasing the volume of the dishes. The ingredients used to increase the volume of rice-based dishes are referred to as \u0026quot;\u003cem\u003ekate\u003c/em\u003e\u0026quot; or \u0026ldquo;\u003cem\u003ekatemono\u003c/em\u003e\u0026rdquo;. Historically, these were regarded as famine foods(62). The use of species to increase the volume of rice dishes has been similarly reported, for example, the use of the roots of \u003cem\u003eDioscorea bulbifera\u003c/em\u003e in Thailand(19).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFig. 4 Botanical families with the greatest number of species used for the four major food categories. Percentages of botanical families for (a) vegetables (n=146), (b) dessert fruit (n=78), (c) beverages (n=23), and (d) snacks (n=19)\u003c/p\u003e\n\u003cp\u003eThe most common parts used as vegetables were shoots (61 species), which represent nearly half of the species used as vegetables. This was followed by leaves, unspecified aerial parts (33 species), roots (19) and stems (18). Five species of inflorescences were used as vegetables, with \u003cem\u003ePetasites japonicus\u003c/em\u003e being the most commonly consumed throughout the country. The young flower buds are called \u003cem\u003efukinoto\u003c/em\u003e and are consumed in a variety of ways, including as tempura and cooked with miso. The \u003cem\u003efukinoto\u003c/em\u003e was described as a plant that sprouts first in spring and heralds the arrival of spring at several study sites. Other species whose inflorescences are eaten as vegetables include \u003cem\u003eWisteria\u0026nbsp;\u003c/em\u003espp. and \u003cem\u003eErythronium japonicum\u003c/em\u003e. Propagules of 3 species were used as vegetables during autumn. The propagules are called \u003cem\u003emukago\u003c/em\u003e in Japanese, and two species of \u003cem\u003emukago\u003c/em\u003e yams (\u003cem\u003eDioscorea japonica\u003c/em\u003e and \u003cem\u003eD. polystachya\u003c/em\u003e) are commonly eaten. They were eaten by roasting, boiling or as the \u003cem\u003ekate\u003c/em\u003e ingredients as mentioned previously. The propagules of \u003cem\u003eElatostema involucratum\u003c/em\u003e were also eaten, although this was limited to one report. The leaves of these plants were removed, and only the stems with propagules were marinated in miso paste. They continue to be consumed in modern times in the northern part of Japan, such as Akita Prefecture, and are popular due to their rarity.\u003c/p\u003e\n\u003cp\u003eThese results are consistent with those reported in previous studies, such as those in Thailand, Laos, and China, which demonstrated a preference for shoots and leaves(15-17, 19), also known as herbophilia(6). Additionally, ferns such as \u003cem\u003ePteridium latiusculum\u003c/em\u003e, \u003cem\u003eOsmunda japonica\u003c/em\u003e, and \u003cem\u003eOncoclea struthiopteris\u003c/em\u003e are also commonly consumed as vegetables in Japan. On the other hand, prior research on the use of wild species in South Korea has indicated that cooked shoots of species from Asteraceae are more frequently consumed as vegetables during the spring season (63-65). Therefore, the findings of the present study were consistent with those previously reported from Korea in terms of the plant taxa and part used, the preparation status (i.e., cooked or uncooked), and the season of use (i.e., spring).\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eDessert Fruits\u003c/h2\u003e\n\u003cp\u003eIn total, there were 78 species of wild plants eaten as dessert fruits, of which almost one-third of the dessert fruits belonged to the Rosaceae family, followed by Ericaceae (8), Elaeagnaceae (4), and Rhamnaceae (4) (Fig. 4(b)). The most popular wild fruit was \u003cem\u003eAkebia\u003c/em\u003e spp. (\u003cem\u003eA. squinata\u003c/em\u003e and \u003cem\u003eA. trifoliata\u003c/em\u003e). Their arils were eaten raw sometime as children\u0026rsquo;s snacks. Other parts of \u003cem\u003eAkebia\u003c/em\u003e are also consumed as vegetables and tea. The stems and buds were consumed as tea and vegetables, while the pericarps were eaten as vegetables in various ways, such as pickled and stir-fried. The second most popular wild fruit was \u003cem\u003eRubus\u003c/em\u003e spp. As mentioned in the Methods section, wild strawberries are called \u003cem\u003enoichigo\u0026nbsp;\u003c/em\u003e(wild strawberry) or \u003cem\u003ekiichigo\u003c/em\u003e (tree strawberry), and there are more than 30 wild \u003cem\u003eRubus\u003c/em\u003e species in Japan, including \u003cem\u003eR. crataegifolius\u003c/em\u003e, \u003cem\u003eR. parvifolius\u003c/em\u003e and \u003cem\u003eR. sieboldii\u003c/em\u003e. Their infructescences were eaten raw, in many cases as children\u0026rsquo;s snacks. However, boiled or pickled fruits are also eaten in some cases. For example, fruits of \u003cem\u003ePyrus pyrifolia\u003c/em\u003e are eaten mostly raw, but in some areas, they are boiled for consumption. There was also a record of fruits of \u003cem\u003eVaccinium oxycoccos\u0026nbsp;\u003c/em\u003ebeing pickled in salt, then sprinkled with sugar and eaten with green tea in the mountainous Oze area in Gunma Prefecture.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eBeverages\u003c/h2\u003e\n\u003cp\u003eThere were 23 species used as beverages, including both alcoholic and nonalcoholic drinks (Fig. 4 (c)). Alcoholic beverages were of two types: so-called wines, in which infructescences were naturally fermented, and so-called liqueurs, in which plants were soaked in spirits such as \u003cem\u003eshochu\u003c/em\u003e. Nonalcoholic beverages included tea and juice.\u003c/p\u003e\n\u003cp\u003eSixteen species were used as teas, including five species of Fabaceae, namely, \u003cem\u003eVicia bungei\u0026nbsp;\u003c/em\u003eand \u003cem\u003eWisteria\u003c/em\u003e spp. Most of them used their aerial parts, such as leaves, shoots and inflorescence, and were processed by steaming, drying or roasting. On the other hand, one study reported that roots were used as tea. The roots of \u003cem\u003eImperata cylindrica\u003c/em\u003e were sun-dried and drunk as tea, which was favored by elderly people because it improves the production of urine.\u003c/p\u003e\n\u003cp\u003eFollowing the tea, seven species were used to make liqueces. Most of them use infructescences for liqueurs, except for the species\u003cem\u003e\u0026nbsp;Geranium thunbergii,\u003c/em\u003e whose aerial parts were used\u003ca href=\"#_ftn1\" name=\"_ftnref1\" title=\"\"\u003e\u003c/a\u003e\u003csup\u003e1\u003c/sup\u003e. In contrast, only two species (\u003cem\u003eVitis coignetiae\u003c/em\u003e and \u003cem\u003eV. ficifolia\u003c/em\u003e) were used for making wine. Although the berries were used, the method of making wine exhibited slight regional variations, including the collection of berries when they were ripe or after frost or fermentation with or without sugar.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSnacks\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSnacks are plants eaten between meals, excluding dessert fruits and nuts. In total, 19 species were eaten as snacks, including 4 from Poaceae and three each from Ericaceae and Polygonaceae (Fig. 4 (d)). Many of these snacks were eaten raw as children\u0026apos;s snacks. Many were aerial parts, such as shoots, leaves, stems, and flowers, and there were many records of children foraging freely for snacks in the fields and mountains.\u003c/p\u003e\n\u003cp\u003eAmong the snacks, many sour-tasting plants, such as \u003cem\u003eRumex japonicus\u003c/em\u003e and Japanese knotweed (\u003cem\u003eReynoutria japonica\u003c/em\u003e), were eaten raw. They were sometimes eaten with salt on the spot of foraging. Similarly, children eating these acidic-tasting species have been documented in various European countries, including Spain, Sweden, and Poland(8). There were other snacks that have a sweet taste, such as common grass (\u003cem\u003eImperata cylindrica\u003c/em\u003e), whose young ears and other parts were eaten; nectar from some flowers, such as Azalea; and the roots of \u003cem\u003eCinnamomum sieboldii\u003c/em\u003e. Another interesting finding was that leaf galls of \u003cem\u003eRhododendron\u003c/em\u003e spp. (Azalea gall) were eaten raw as snacks, although this finding was limited to three reports. Azalea leaf galls are caused by fungi, including \u003cem\u003eExobasidium\u003c/em\u003e spp. (66), and locally called \u003cem\u003emochi\u003c/em\u003e (rice cake), the disease is called \u003cem\u003emochi\u003c/em\u003e disease. There have been reports of the use of plant galls in other parts of the world, including as folk medicine and for dyeing(67-69). However, reports of Azalea galls as snacks for children appear to be limited to the present study.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStarches\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe 17 species used as starches were largely divided into those that use roots and those that use seeds. The most common family associated with the roots was Liliaceae (4 species), whereas the most common family associated with the seeds was Fagaceae (4 species), which included 3\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cem\u003eQuercus\u003c/em\u003e spp. (\u003cem\u003eQ. acuta\u003c/em\u003e, \u003cem\u003eQ. gilva,\u003c/em\u003e and\u003cem\u003e\u0026nbsp;Q. glauca\u003c/em\u003e). Various techniques were used to extract starches, such as pounding, grating, soaking in water and strain. These extracted starches were used for making rice cakes, noodles, porridges and jelly type foods.\u003c/p\u003e\n\u003cp\u003eTwo toxic plants (\u003cem\u003eLycoris radiata\u003c/em\u003e and \u003cem\u003eCycas revolute\u003c/em\u003e) were included in this category. Although it was limited to only one report, bulbs of \u003cem\u003eL. radiata\u003c/em\u003e were used for making starches. \u003cem\u003eL. radiata\u003c/em\u003e contains lycorine, which is commonly found in the Amaryllidaceae (70). People were aware of this toxicity; therefore, they took a long time to remove the poison by being exposed to water repeatedly over a long period of time. Another poisonous plant was \u003cem\u003eCycas revolute\u003c/em\u003e. Their seeds and stems were used for extracting starches, particularly from the Ryukyu and Amami Islands of southern Japan. Like the use of \u003cem\u003eL. radiata\u003c/em\u003e, to remove toxins, including cycasin, the plant parts used were exposed to water repeatedly over time. In some areas, people ferment them, and some report the infestation of maggots as a sign of detoxification.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eBinders\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePlants used to combine food ingredients are categorized as binders or \u0026ldquo;\u003cem\u003etsunagi\u003c/em\u003e\u0026rdquo; in Japanese. They were mostly used for rice cakes (\u003cem\u003emochi\u0026nbsp;\u003c/em\u003eand \u003cem\u003edango\u003c/em\u003e) and sometimes for \u003cem\u003esoba\u003c/em\u003e (buckwheat noodles). Among the 13 species used in this category, there were 8 species of Asteraceae, including two \u003cem\u003eArtemisia\u0026nbsp;\u003c/em\u003espp. (\u003cem\u003eA. princeps\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;indica\u003c/em\u003e) and 3 \u003cem\u003eSynurus\u003c/em\u003e spp. (\u003cem\u003eS. pungens, deltoides\u003c/em\u003e and \u003cem\u003eexcelsus\u003c/em\u003e). Young leaves and shoots of these species were boiled and added to food, and their trichomes and fibers contribute to the binding of other ingredients. In addition, some binder plants, such as \u003cem\u003eArtemisia\u003c/em\u003e spp. and \u003cem\u003ePseudognaphalium affine,\u003c/em\u003e also contribute to the aroma or color of the prepared food. Although it was limited to two reports, the bark and cortex of red pine (\u003cem\u003ePinus densiflora\u003c/em\u003e) were used to make rice cakes called \u003cem\u003ematsukawa mochi\u003c/em\u003e (pine skin rice cakes). Compared with other parts of the world, \u003cem\u003eA. spp\u003c/em\u003e. and other species have been used as ingredients for mochi in China and Korea (71-73). Some research has also indicated that the use of \u003cem\u003eP. affine\u003c/em\u003e for mochi in Japan was originally introduced from China and was later replaced by \u003cem\u003ethe use of A.\u0026nbsp;\u003c/em\u003espp. as an alternative to P. affine (74).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eNuts\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAmong the 13 species consumed as nuts, Fagaceae\u0026nbsp;presented\u0026nbsp;the greatest number of species (6 species). Although most species\u0026nbsp;were tree plants,\u0026nbsp;the seeds of 3 species of aquatic plants\u0026nbsp;were\u0026nbsp;also eaten as nuts (\u003cem\u003eTrapa natans\u003c/em\u003e var.\u003cem\u003e\u0026nbsp;Bispinosa, Nelumbo nucifera,\u0026nbsp;\u003c/em\u003eand \u003cem\u003eEuryale ferox\u003c/em\u003e\u003cem\u003ee\u003c/em\u003e). These nuts were eaten raw, roasted, or boiled and sometimes as snacks.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eOils/fats\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThere were only 3 species used for oils/fats (\u003cem\u003eTerreya nucifera\u003c/em\u003e, \u003cem\u003eCamellia\u0026nbsp;\u003c/em\u003e\u003cem\u003ejaponica\u003c/em\u003e, and\u003cem\u003e\u0026nbsp;C.\u0026nbsp;\u003c/em\u003e\u003cem\u003eSasanqua\u003c/em\u003e). Oils were extracted from seeds of these species. Among the three species, \u003cem\u003eC. japonica\u003c/em\u003e was the most frequently reported, although its use was limited to only 6 reports in western Japan, particularly in the Kyushu region. The other two species were reported only once each. These oils were considered valuable and were sometimes sold and given as gifts.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eOther Food Types\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis category included the １3 species used for coloring, flavoring, fermented foods, or algae jelly making. For coloring and flavoring, for example, \u003cem\u003eGardenia jasminoides\u0026nbsp;\u003c/em\u003ewas used to add yellow color to food, including rice cakes and pickled vegetables. Another example is chick weeds (\u003cem\u003eStellaria media\u003c/em\u003e). In addition to their use as vegetables, they were also used for coloring and flavoring. In the Nasu area of Tochigi Prefecture, their aerial parts were used along with \u003cem\u003eshiso\u003c/em\u003e leaves (\u003cem\u003ePerilla frutescens\u003c/em\u003e) to provide sourness and better color when pickling \u003cem\u003ewith\u003c/em\u003e plums (\u003cem\u003ePrunus mume\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003eThere were 8 species used for fermented foods, and their reports were all regionally specific. Some species (\u003cem\u003eHumulus cordifolius\u003c/em\u003e and \u003cem\u003eEquisetum arvense\u003c/em\u003e) wereadded when alcoholic beverages such as rice wine were made since they were considered to accelerate fermentation in the Tohoku region. In Nagano Prefecture, juices made from infructescences of 3 species (\u003cem\u003eMalus toringo\u003c/em\u003e, \u003cem\u003ePyrus pyrifolia\u003c/em\u003e,\u003cem\u003e\u0026nbsp;and Cycas revoluta\u003c/em\u003e) were used for unsalted lactic-fermented pickles called \u003cem\u003esunki\u003c/em\u003e. These compounds are called \u0026ldquo;sources\u0026rdquo; or \u0026ldquo;essences\u0026rdquo; of \u003cem\u003esunki\u0026nbsp;\u003c/em\u003epickles and are believed to contribute to pH control and the production of lactic acid, etc. (75). As mentioned previously, the use of \u003cem\u003eCycas revolute\u003c/em\u003e was limited to only southern islands. On Amami islands, seeds of \u003cem\u003eC . revolute\u0026nbsp;\u003c/em\u003ewere also used to make \u003cem\u003ekoji\u003c/em\u003e molds (fermentation starters) for making fermented soy bean paste, \u003cem\u003emiso\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eTwo species (\u003cem\u003eOxalis corniculata\u003c/em\u003e and \u003cem\u003eVitis coignetiae\u003c/em\u003e) were used to make \u003cem\u003etokoroten\u003c/em\u003e, jelly noodles made from red algae, including \u003cem\u003eGelidium amansii\u003c/em\u003e. The use of \u003cem\u003eO. corniculata\u003c/em\u003e was reported only once in Ishikawa Prefecture, while there were 3 reports of the use of \u003cem\u003eV. coignatiae\u003c/em\u003e in the Tohoku and Hokkaido regions. The parts used and how to use them differ from each other. Aerial parts of \u003cem\u003eO. corniculate\u0026nbsp;\u003c/em\u003eor unripen fruits of \u003cem\u003eV. coignetiae\u003c/em\u003e were used when boiling red algae instead of vinegar. These compounds are considered to help extract agarose from red algae(76). Sour-tasting saps from vines of \u003cem\u003eV. coignetiae\u0026nbsp;\u003c/em\u003ewere also used after red algae were boiled, but they were used to curdle the agar jelly. Thus, these sour plants were used to extract or solidify the jelly.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eToxic\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;Plants\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThere were several reports of the use of toxic plants, although the type and degree of toxicity vary. For example, species containing carcinogens were eaten. As mentioned, bracken (\u003cem\u003ePteridium latiusculum\u003c/em\u003e) is commonly eaten throughout Japan, with shoots being eaten as vegetables and bracken starch extracted from the roots. Bracken contains a carcinogen called ptaquiloside (77). Another plant that contains carcinogens is \u003cem\u003eCycas revolute.\u003c/em\u003e Stems and seeds of \u003cem\u003eC. revolute\u003c/em\u003e were used to make starch and fermentation starters; however, C. revolute contains a carcinogen called cycasin, which is found in other cycads(78).\u003c/p\u003e\n\u003cp\u003eAlkaloid-containing species were also consumed. For example, rhizomes of\u003cem\u003e\u0026nbsp;Dioscorea\u003c/em\u003e spp. (specifically \u003cem\u003eD. tokoro\u003c/em\u003e) were boiled and consumed. Their rhizomes contain dioscin, a type of saponin that is commonly found in \u003cem\u003eDioscorea\u003c/em\u003e spp. (79). It was initially reported to have a very bitter taste but became palatable once people became accustomed to it in Saga Prefecture. In two of the five reports of \u003cem\u003eD. tokoro\u0026nbsp;\u003c/em\u003eas food, they were also used as an offering. The use as offerings to the gods has been reported in several places in Japan(36).\u0026nbsp;Another alkaloid-containing species consumed was\u003cem\u003e\u0026nbsp;Lycoris radiata\u003c/em\u003e for starch from its bulbs, as mentioned previously.\u003c/p\u003e\n\u003cp\u003eOne of the most toxic plants eaten is the lacquer tree (\u003cem\u003eToxicodendron vernicifluum\u003c/em\u003e), which widely grows in China, Korea and Japan. In Asia, sap is used as a lacquer, but it contains an allergen called urushiol(80). In Miyagi and Ibaraki prefectures, their shoots were consumed as vegetables. In Miyagi, older individuals liked to consume shoots because of their soft texture, whereas in Ibaraki, they were regarded as tasty but potentially irritating food, with the decision to consume them dependent on the individual. It has been reported that individuals engaged in lacquer production, such as artisans, adhere to the custom of eating lacquer plants as a means of reducing their susceptibility to lacquer rash in Japan(81). However, the two regions reported in this study were not lacquer-producing areas and therefore cannot be relevant to this case of lacquer eating for immunity. On the other hand, the consumption of leafy shoots of \u003cem\u003eT. vernicifluum\u003c/em\u003e as vegetables has been reported among ethnic minority groups in China, such as Lemo and Dulong in Yunnan Province and Yi in Sichuan Province(22, 82, 83). To understand the food use of lacquers in Japan, it is necessary to consider the food use of lacquers in the wider context of East Asia.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eProcessing Techniques\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe majority of plants, particularly vegetables, were consumed primarily in cooked form. However, a range of other processing techniques were also employed.\u003c/p\u003e\n\u003cp\u003eIn particular, almost 300 reports of \u003cem\u003eakunuki,\u003c/em\u003e a food processing technique used to remove \u003cem\u003eaku\u003c/em\u003e, which are unpalatable and undesirable substances or tastes such as harshness and astringency, existed. The \u003cem\u003eakunuki\u0026nbsp;\u003c/em\u003emethods include the use of various ingredients, such as ash, rice water (water used to wash rice), rice bran, baking soda, vinegar, and camellia leaves; prolonged soaking in water; prolonged burial in soil; fermentation; and a combination of these methods. Some of these techniques are capable of removing not only unpleasant tastes but also toxic substances from wild species. The most common \u003cem\u003eakunuki\u003c/em\u003e method was the use of ashes (179 reports). In many cases, the plant parts were soaked in water with ash for a long time or boiled. Among the species that were pretreated with ashes, \u003cem\u003ewarabi\u003c/em\u003e bracken (\u003cem\u003ePteridium latiusculum\u003c/em\u003e) was reported most frequently (80 reports), followed by \u003cem\u003eArtemisia princeps\u003c/em\u003e (31), \u003cem\u003eOsmunda japonica\u003c/em\u003e (28) and \u003cem\u003eAesculus turbinate\u0026nbsp;\u003c/em\u003e(22). These reports included detailed descriptions of the specific types of ash to be used, for example, ash from stoves and hearths, rice straw ash and wood ash. There were several reports of the use of specific types of ash, particularly in the pretreatment of \u003cem\u003eAesculus turbinate\u003c/em\u003e seeds when they weremixed with glutinous rice to produce rice cakes (\u003cem\u003etochi mochi\u003c/em\u003e). Among the 22 reports, more than half described specific types of ashes to be used, including those of hardwoods, mainly \u003cem\u003eQuercus\u003c/em\u003e spp. The use of ash, which is alkaline, for plant detoxification is a well-known traditional method involving pH control (84). The reason for the use of hardwood ashes, such as those derived from \u003cem\u003eQuercus\u003c/em\u003e spp., is likely due to the pronounced alkalinity of these ashes, which has a greater impact on detoxification(85).\u003c/p\u003e\n\u003cp\u003eThe second most common method used for \u003cem\u003eakunuki\u003c/em\u003e was the use of rice bran (58 reports). The plant parts were boiled in water with rice bran or, alternatively, left over water after rising rice was used. This method is commonly employed for the preparation of a variety of bamboo shoots. The use of rice bran is likely to facilitate the adsorption of undesired substances(84). However, in other Asian countries where bamboo shoots are widely consumed, boiling in water or saltwater is the predominant pretreatment method (86). A previous study suggested that the method of using rice bran is a relatively recent phenomenon that emerged after the late Meiji period (1868\u0026ndash;1912) (87).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMedicinal plants\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePrevious studies have demonstrated a blurred boundary between food and medicine in the use of wild plants(88, 89). This study also identified 28 plant species that were used for food, medicinal and health-promoting purposes (Table 2). These plants were consumed for the treatment of various health conditions, including detoxication, deworming, cough, fever, tinnitus, and dizziness. Some of them were consumed specifically by infants, children, pregnant women, postpartum women and elderly individuals. The most frequently occurring species were those belonging to the Asteraceae family (5 species), followed by those of Lamiaceae (3 species). The species that were consumed as medicines included those that were commonly eaten as food throughout the country. For example, there were three reports of juice extracted from buds or roots of \u003cem\u003ePetasites japonicus\u003c/em\u003e being given to newborns to detoxify the embryotoxic (\u003cem\u003etaidoku\u003c/em\u003e)(90, 91)\u003ca href=\"#_ftn2\" name=\"_ftnref2\" title=\"\"\u003e\u003c/a\u003e\u003csup\u003e2\u003c/sup\u003e. Another example was \u003cem\u003eOsmunda japonica.\u003c/em\u003e In the Tohoku region, they are believed to be beneficial for consumption after childbirth, with the aim of improving blood circulation or cleansing blood. Conversely, in Niigata Prefecture, eating them after birth was prohibited because they believed that they could cause blood to become \u0026ldquo;rough\u0026rdquo;. There were also six species, including \u003cem\u003eArtemisia princeps,\u003c/em\u003e that were consumed as herbal teas, which are generally regarded as beneficial for the body.\u003c/p\u003e\n\u003cp\u003eTable 2. A list of species consumed as food as well as for medicinal and health-related purposes\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"632\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003eScientific name\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eFamily\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eParts used and preparations\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003eFunctions\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eActinidia polygama\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eActinidiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eseeds\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003estomach\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eTetragonia tetragonoides\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAizoaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eleaves\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003egastrointestinal\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eOenanthe javanica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eApiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eshoots, boiled and mixed with miso or soy-sauce\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edetoxication\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eArtemisia indica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003ejuice from aerial parts\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003efever\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eArtemisia princeps\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eherbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eAster yomena\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eshoots, boiled and mixed with miso or soy-sauce\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edetoxication\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eCircium\u0026nbsp;\u003c/em\u003espp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eleaves, shoots\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eCrepidiastrum lanceolatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003ebrew herbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003efever\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eFarfugium japonicum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003estems, boiled or salted\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edetoxication\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePetasites japonicus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003ejuice roots\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edetoxication particularly for newborn(roots)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePseudognaphalium affine\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eAsteraceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003ebrew herbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003ecough\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePlatycodon grandiflorus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eCampanulaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eroots, pickled\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003ecough\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eElaeagnus umbellata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eElaeagnaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eherbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003etinnitus, deafness\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePueraria montana\u0026nbsp;\u003c/em\u003evar. \u003cem\u003elobata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eFabaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003edrink hot water mixed with starch extracted from roots\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003efood for the sick\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eGeranium thunbergii\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eGeraniaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eClerodendrum trichotomum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eLamiaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eshoots, cooked\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edeworming, prevent summer fatigue\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eErythronium japonicum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eLiliaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003edrink hot water mixed with starch extracted from roots\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003efood for the sick\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eOsmunda japonica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eOsmundaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eshoots, cooked\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003ecleansings of blood during childbirth\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePlantago asiatica\u003c/em\u003e var. \u003cem\u003easiatica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003ePlantaginaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eshoots, boiled and mixed with miso or soy-sauce, herbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edetoxication\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eImperata cylindrica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003ePoaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eroots, brewed as herbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003epromoting urination\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eHovenia dulcis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eRhamnaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003einfructescences, dried\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePyrus pyrifolia\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eRosaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eriped infructescences, raw\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003ecough, prevent asthma\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003ePhellodendron amurense\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eRutaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003edebarked stems, brewed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003estomachach\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eHouttuynia cordata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eSaururaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eherbal tea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eAlkekengi officinarum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eSolanaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003einfructescences, raw\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003en/a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eTorreya nucifera\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eTaxaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eseeds\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edeworming\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eCamellia japonica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eTheaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003eoils from seeds\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edeworming for children\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"26.856240126382307%\"\u003e\n \u003cp\u003e\u003cem\u003eVitis coignetiae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.429699842022117%\"\u003e\n \u003cp\u003eVitaceae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.859399684044234%\"\u003e\n \u003cp\u003einfructescences, wine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.85466034755134%\"\u003e\n \u003cp\u003edizziness, good for health in general and for pregnant woman\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eMale and Female Plants\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThere were 5 reports on the uses of male and female plants of the 4 species. In Gifu Prefecture, stems of \u003cem\u003eReynoutria japonica\u003c/em\u003e were eaten raw as snacks with salt. The stems of the female plants were preferred because they were thicker, softer, and tasted sour, whereas the stems of the male plants were thinner and harder. Additionally, some reports indicate that plant sex determines whether a plant is used. For example, in Amami Ōshima Island of Kagoshima Prefecture, stems of \u003cem\u003eCycas revol\u003c/em\u003e\u003cem\u003euta\u003c/em\u003e were processed and used as starches. Only male plants were selected for starch extraction, as male plants were considered to store more starch. Another example involves approximately 2 species of Liliaceae (\u003cem\u003eCardiocrinum cordatum\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;Lilium auratum var. auratum\u003c/em\u003e), where the roots of the female plants were only selected for starch extraction. \u003cem\u003eReynoutria japonica\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;Cycas revoluta\u003c/em\u003e are dioecious plant species, whereas\u003cem\u003e\u0026nbsp;Lilium auratum var. auratum\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;Cardiocrinum cordatum\u003c/em\u003e are not. Thus, preferences for texture and flavor and whether to use plants were determined according to the sex of the plants. However, how male and female plants were locally identified for these 4 species was not described in the literature. As there are only a few studies on traditional knowledge of plant dioecy and the folk concepts of plant sex, further research is needed on the local perceptions of plant sex, as suggested by previous studies(92, 93).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTaste and flavor\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThere were a number of descriptions pertaining to the taste and flavor of wild species. These can be broadly classified into those pertaining to flavor, texture, and taste. Flavors were described as being of good quality, specific to wild species, in the spring, with all of them being perceived as positive. For example, in Akita, \u003cem\u003eHasteola robusta\u003c/em\u003e was carefully cooked as \u003cem\u003etempura\u003c/em\u003e to retain its distinctive aroma, which is considered important. There were several reports about texture, including the sliminess of \u003cem\u003eElatostema involucratum\u003c/em\u003e, the crispness of \u003cem\u003eAstilbe rubra\u003c/em\u003e, and the stickiness and gooeyness of \u003cem\u003eL. auratum\u0026nbsp;\u003c/em\u003evar.\u003cem\u003e\u0026nbsp;auratum\u003c/em\u003e. These attributes were all regarded as positive. In terms of taste, wild species were often described as having a distinctive and slightly bitter taste, which wasperceived as good. This was exemplified in the preparation of seeds of \u003cem\u003eAesculus turbinata\u003c/em\u003e and bulbs of\u003cem\u003e\u0026nbsp;Lilium auratum\u003c/em\u003e var. \u003cem\u003eauratum\u003c/em\u003e, where some described taking care not to lose all of the bitterness. There were also nine reports that the taste of wild species was better than that of cultivated or purchased species or those grown in proximity to their residences. Examples of these include \u003cem\u003eudo\u003c/em\u003e (\u003cem\u003eAralia cordata\u003c/em\u003e), butterbur (\u003cem\u003ePetasites japonicus\u003c/em\u003e),\u003cem\u003e\u0026nbsp;\u003c/em\u003ehorse\u0026nbsp;chestnut\u0026nbsp;(\u003cem\u003eCastanea crenata\u003c/em\u003e)\u003cem\u003e\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;\u003c/em\u003eyam (\u003cem\u003eDioscorea japonica\u003c/em\u003e). Several previous studies have shown that wild vegetables were appreciated and preferred because of their tastes, for example, in southern Italy, Thailand, Himalaya and Japan (39, 94-97). Wild food plants were considered not only a source of nutrition or a means of sustenance but also a source of enjoyment through their unique tastes and flavors in Japan.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eUse of\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003erare species\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThere\u0026nbsp;were\u0026nbsp;few reports on the use of threatened species according to their current conservation status.\u0026nbsp;Among the\u0026nbsp;2163 species of vascular plants listed\u0026nbsp;on the\u0026nbsp;2020 Japanese Red List(98), the following four species were used approximately 1930:\u003cem\u003e\u0026nbsp;Platycodon grandifloras\u003c/em\u003e,\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eprickly waterlily (\u003cem\u003eEuryale ferox\u003c/em\u003e), and bird\u0026rsquo;s nest fern (\u003cem\u003eAsplenium antiquum\u003c/em\u003e),\u003cem\u003e\u0026nbsp;\u003c/em\u003ewhich are listed as endangered, and \u003cem\u003eCinnamomum sieboldii\u003c/em\u003e, which is listed as near threatened. The roots of \u003cem\u003eP. grandifloras\u003c/em\u003e were pickled in plum vinegar and were traditionally consumed in Hiroshima Prefecture. They are also used as a cough medicine, although their taste was not very tasty. There were 2 reports about the use of \u003cem\u003eEuryale ferox\u003c/em\u003e. The cooked stems were eaten as\u0026nbsp;vegetables, whereas\u0026nbsp;the roasted or boiled seeds were eaten as snacks. Additionally, dried and\u0026nbsp;ground\u0026nbsp;seeds were used as starches to make dumplings (\u003cem\u003edango\u003c/em\u003e). \u003cem\u003eAsplenium antiquum\u003c/em\u003e was reported once in Ishigaki city in Okinawa, where the shoots were used as vegetables in simmered or dressed dishes. \u003cem\u003eCinnamomum sieboldii\u003c/em\u003e was reported twice. The roots were eaten raw as snacks for the children.\u0026nbsp;\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThis study examined Japan's largest food culture database and reviewed the food use of wild plants in the early Showa period (1926\u0026ndash;1989). At least 268 taxa of plants belonging to 88 families were used to make food and beverages in Japan. Asteraceae was the most frequently reported and most species-rich family. The majority of species was documented in only a limited number of reports, and the number of species used at the national level was limited. In other words, each region and locality had its own distinctive culinary traditions involving wild food species. A comparison with those of previous studies on wild species used in other Asian regions revealed a similar preference for vegetables, namely, shoots and leaves. However, notably, in Japan, ferns were also consumed in considerable quantities. Furthermore, there were some other similarities with other Asian regions, including folk uses of plants such as binder plants and \u003cem\u003ekate\u003c/em\u003e plants and the consumption of toxic plants (lacquers). Conversely, further elucidation is clearly needed with respect to certain matters, including the perception of plant sex, the relationship between the consumption of lacquer and immunity, and the relationships between diverse ash components and detoxification techniques. It is therefore evident that further ethnobotanical research is needed, both in the form of an examination of ethnographic records and in the field, to gain a deeper understanding of the use of wild food plants in Japan.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eNot Applicable\u003c/p\u003e\n\u003ch2\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eNot Applicable\u003c/p\u003e\n\u003ch2\u003eCompeting interests\u003c/h2\u003e\n\u003cp\u003eThe authors declare that they have no competing interests\u003c/p\u003e\n\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eLotte Foundation\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eThe author confirms sole responsibility from conceptualizing and designing study , data collection, analysis and interpretation of results, and to manuscript preparation.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eThis research was supported by JSPS KAKENHI Grant Numbers JP21H03685, JP21K01086, and Lotte Foundation.\u003c/p\u003e\n\u003ch2\u003eAvailability of data and materials\u003c/h2\u003e\n\u003cp\u003eI confirm that the data supporting the findings of this study are available within the article and its supplementary material.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eB\u0026eacute;langer J, Pilling D. 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A comparative study on wild food plant consumption in twenty-one local communities in Italy. J Ethnobiol Ethnomed. 2007;3:22.\u003c/li\u003e\n\u003cli\u003eSomnasang P, Moreno-Black G. Knowing, gathering and eating: knowledge and attitudes about wild food in an Isan village in Northeastern Thailand. Journal of Ethnobiology. 2000;20(2):197-216.\u003c/li\u003e\n\u003cli\u003eThakur D, Sharma A, Uniyal SK. Why they eat, what they eat: patterns of wild edible plants consumption in a tribal area of Western Himalaya. Journal of ethnobiology and ethnomedicine. 2017;13:1-12.\u003c/li\u003e\n\u003cli\u003eSugiura T, Kishimoto K. On the liking of wild vegetables and mushrooms by consumers. Journal of the Japanese Forestry Society. 1989;71(1):31-8.\u003c/li\u003e\n\u003cli\u003eEnvironment JMot. Japanese Red List 2020. In: Environment JMot, editor. 2020.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Footnotes","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003e \u003cem\u003eG. thunbergii\u003c/em\u003e was a common folk medicine in Japan. A number of reports in the collection also showed that they were used as a medicine. As explained in the Methods, plants that were only used as medicines were not included in this study. This particular report was included since \u003cem\u003eG. thunbergii\u003c/em\u003e was described as a wild plant used to make herbal liquors which had both food and medicinal purposes.\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003e At the time, it was believed that \u003cem\u003etaidoku\u003c/em\u003e could cause neonatal illnesses such as boil and rash, and plants and medicines were given to infants shortly after birth. This theory of embryonic poisoning is thought to have originate in China.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Wild food plants, food, wild species, traditional knowledge, Japan, Asia","lastPublishedDoi":"10.21203/rs.3.rs-4803162/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4803162/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground\u003c/h2\u003e \u003cp\u003eJapan, which has a diverse climate, is home to 8118 species of wild vascular plants, with more than 1,000 of these species considered edible. However, there are fewer studies on the use of wild food plants in Japan than in other parts of the world. This research aims to provide an ethnobotanical review of the traditional food use of wild plants in Japan. It draws upon the largest database of traditional Japanese foodways, with a specific focus on the period approximately 1930. This occurred prior to Japan\u0026rsquo;s rapid economic growth, which resulted in significant changes in the use of wild plants.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eThe use of wild food plants in Japan was reviewed by studying the collection of Japanese foodways (\u003cem\u003eNihon no shokuseikatsu zenshu\u003c/em\u003e). The review extracted the local and common names, use locations, uses, processing and cooking methods, habitats and other relevant details pertaining to the reported use of wild food plants. This information was subsequently analyzed via Excel.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eA total of 268 taxa belonging to 87 families used in Japan approximately 1930 were documented. Asteraceae was the most frequently reported family as well as the most species-rich family. In terms of use, vegetables composed the largest group, followed by dessert fruits. The majority of species have been documented in only a limited number of reports. The review revealed local traditional uses of plants, including the use of plants as binders for making rice cake, the selection of plants according to their sex, and techniques for detoxifying toxic plants.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e \u003cp\u003eA comparison with those of previous studies on wild species used in other Asian regions revealed a similar preference for vegetables. However, notably, in Japan, ferns are also consumed in considerable quantities. Conversely, further elucidation is clearly needed with respect to certain matters, including the perception of plant sex and the relationships between diverse ash components and detoxification techniques. It is evident that further ethnobotanical research is needed, both in the form of an examination of ethnographic records and in the field, to gain a deeper understanding of the use of wild food plants in Japan.\u003c/p\u003e","manuscriptTitle":"Ethnobotanical review of traditional use of wild food plants in Japan","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-08-27 09:45:14","doi":"10.21203/rs.3.rs-4803162/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-08-10T22:57:51+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-08-10T22:39:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-08-07T08:09:19+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"164489523909801140654706031110113065539","date":"2024-07-31T08:39:54+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"181375491418148040642805843717580188053","date":"2024-07-31T08:28:50+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-07-31T07:43:40+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-07-31T00:45:57+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-07-31T00:45:29+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Ethnobiology and Ethnomedicine","date":"2024-07-25T16:04:32+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-ethnobiology-and-ethnomedicine","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jeet","sideBox":"Learn more about [Journal of Ethnobiology and Ethnomedicine](http://ethnobiomed.biomedcentral.com/)","snPcode":"13002","submissionUrl":"https://submission.nature.com/new-submission/13002/3","title":"Journal of Ethnobiology and Ethnomedicine","twitterHandle":"@ethnobiomed","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"bbe01008-6aa8-44b4-8428-055ca0ef29ce","owner":[],"postedDate":"August 27th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-11-25T16:04:15+00:00","versionOfRecord":{"articleIdentity":"rs-4803162","link":"https://doi.org/10.1186/s13002-024-00736-2","journal":{"identity":"journal-of-ethnobiology-and-ethnomedicine","isVorOnly":false,"title":"Journal of Ethnobiology and Ethnomedicine"},"publishedOn":"2024-11-21 15:57:06","publishedOnDateReadable":"November 21st, 2024"},"versionCreatedAt":"2024-08-27 09:45:14","video":"","vorDoi":"10.1186/s13002-024-00736-2","vorDoiUrl":"https://doi.org/10.1186/s13002-024-00736-2","workflowStages":[]},"version":"v1","identity":"rs-4803162","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4803162","identity":"rs-4803162","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}