{"id":1166,"date":"2023-05-09T15:03:25","date_gmt":"2023-05-09T06:03:25","guid":{"rendered":"https:\/\/www.nibb.ac.jp\/plantdic\/blog\/?p=1166"},"modified":"2026-04-03T08:26:21","modified_gmt":"2026-04-02T23:26:21","slug":"%e9%a2%a8%e5%aa%92%e3%81%ae%e3%82%b9%e3%83%9a%e3%82%b7%e3%83%a3%e3%83%aa%e3%82%b9%e3%83%88%e3%80%81%e3%83%a4%e3%83%9e%e3%83%88%e3%82%b0%e3%82%b5%e5%b1%9e-specialist-of-wind-pollination-theligonum","status":"publish","type":"post","link":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/?p=1166","title":{"rendered":"\u98a8\u5a92\u306e\u30b9\u30da\u30b7\u30e3\u30ea\u30b9\u30c8\u3001\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e Specialist of wind-pollination Theligonum"},"content":{"rendered":"\n
\"\u30e4\u30de\u30c8\u30b0\u30b5\u3000Theligonum<\/a>
\u56f3\uff11\u3000\u30e4\u30de\u30c8\u30b0\u30b5\u3000Theligonum japonicum<\/em>: (A-C) \u30e4\u30de\u30c8\u30b0\u30b5\u306e\u958b\u82b1\u500b\u4f53\uff08A\uff09\u3001\u82b1\u306e\u62e1\u5927\uff08B)\u3001\u672a\u958b\u82b1\u500b\u4f53\uff08C) \u3002<\/figcaption><\/figure>\n\n\n\n

\u30e4\u30de\u30c8\u30b0\u30b5\u5c5eTheligonum<\/em>\u306f\u3001\uff14\u7a2e\u304c\u77e5\u3089\u308c\u3066\u3044\u308b\u3002\u65e5\u672c\u3068\u4e2d\u56fd\u6771\u90e8\u306b\u30e4\u30de\u30c8\u30b0\u30b5 Theligonum japonicum<\/em> Okubo & Makino (Chen and Funston 2011)\u3001\u53f0\u6e7e\u306bT<\/em>. formosanum<\/em> (Ohwi) Ohwi & T.S. Liu (Chen and Funston 2011)\u3001\u4e2d\u56fd\u4e2d\u897f\u90e8\uff08\u56db\u5ddd\u7701\u3001\u96f2\u5357\u7701\u306a\u3069\uff09\u306bT. macranthum <\/em>Franchet (Chen and Funston 2011)\u3001\u5730\u4e2d\u6d77\u6cbf\u5cb8\u306bT. cynocrambe <\/em>L. (https:\/\/www.gbif.org\/species\/2919295)\u3001\u304c\u5206\u5e03\u3059\u308b\u3002<\/p>\n\n\n\n

Four species of the genus Theligonum<\/em> are known: Theligonum japonicum<\/em> Okubo & Makino (Chen and Funston 2011) in Japan and eastern China, T. formosanum<\/em> (Ohwi) Ohwi & T.S. Liu (Chen and Funston 2011) in Taiwan, T. macranthum<\/em> Franchet (Chen and Funston 2011) in central and western China (Sichuan Prov, and Yunnan Prov.), T. macranthum<\/em> Franchet (Chen and Funston 2011) in central and western China (Sichuan, Yunnan, etc.), T. cynocrambe L. (https:\/\/www.gbif.org\/species\/2919295) on the Mediterranean coast.<\/p>\n\n\n\n

(A-C) T. japonicum<\/em>: Flowering individuals (A), enlarged flowers (B), and non-flowering individuals (C).<\/p>\n\n\n\n

\"\u30a2\u30ab\u30cd\u4e9c\u79d1\u306e\u5206\u5b50\u7cfb\u7d71\u6a39\"<\/a>
\u56f3\uff12\u3000\u30a2\u30ab\u30cd\u4e9c\u79d1\u306e\u5206\u5b50\u7cfb\u7d71\u6a39<\/figcaption><\/figure>\n\n\n\n

\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u306f\u3001\u30a2\u30ab\u30cd\u79d1\u3001\u30a2\u30ab\u30cd\u4e9c\u79d1\u3001\u30e4\u30de\u30c8\u30b0\u30b5\u9023\u306b\u5c5e\u3059\u308b\u3002\u30e4\u30de\u30c8\u30b0\u30b5\u9023\u306b\u306f\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u306e\u307f\u3092\u542b\u3080\u3002 <\/p>\n\n\n\n

The genus Theligonum<\/em> belongs to the family Rubiaceae, subfamily Rubioideae, and section Theligoneae<\/em>. The section Theligoneae<\/em> includes only the genus Theligonum<\/em>.<\/p>\n\n\n\n

\"\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u6804\u990a\u30b7\u30e5\u30fc\u30c8\u3068\u751f\u6b96\u30b7\u30e5\u30fc\u30c8\u3000Vegetative<\/a>
\u56f3\uff13\u3000\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u6804\u990a\u30b7\u30e5\u30fc\u30c8\u3068\u751f\u6b96\u30b7\u30e5\u30fc\u30c8\u3000Vegetative and reproductive shoots of Theligonum japonicum<\/em><\/figcaption><\/figure>\n\n\n\n

\u30e4\u30de\u30c8\u30b0\u30b5\u306f\u96cc\u96c4\u7570\u82b1\u540c\u682amonoecious\u3067\u3001\u540c\u3058\u500b\u4f53\u304c\u96c4\u82b1\u3068\u96cc\u82b1\u3092\u5f62\u6210\u3059\u308b\uff08A\u3001B)\u3002\u6804\u990a\u30b7\u30e5\u30fc\u30c8\u3067\u306f\u4ed6\u306e\u30a2\u30ab\u30cd\u79d1\u306e\u7a2e\u540c\u69d8\u3001\u8449\u306f\u5bfe\u751f\u3059\u308b\uff08C\u3001D)\u3002\u4e00\u65b9\u3001\u82b1\u3092\u4ed8\u3051\u308b\u751f\u6b96\u30b7\u30e5\u30fc\u30c8\u3067\u306f\u3001\u8449\u3068\u5bfe\u751f\u306e\u4f4d\u7f6e\u306b\uff12\u500b\u306e\u96c4\u82b1\u304c\u5f62\u6210\u3055\u308c\u308b\uff08E\u3001F)\u3002\u96c4\u82b1\u306f\u9000\u5316\u6d88\u5931\u3057\u305f\u8449\u306e\u8449\u814b\u306b\u5f62\u6210\u3055\u308c\u3066\u3044\u308b\u3068\u8003\u3048\u3089\u308c\u308b\u3002<\/p>\n\n\n\n

T. japonicum<\/em> is monoecious, with the same individual forming male and female flowers (A, B). In the vegetative shoot, the leaves are opposite (C, D), as in other Rubiaceae species. On the other hand, in the flowering reproductive shoot, two male flowers are formed in the opposite position to the leaf (E, F). The male flowers are hypothesized to be formed in the axils of the degenerated and lost leaves.<\/p>\n\n\n\n

\"\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u96c4\u82b1<\/a>
\u56f3\uff14\u3000\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u96c4\u82b1 Male flowers of Theligonum japanicum<\/em><\/figcaption><\/figure>\n\n\n\n

\u6210\u719f\u3057\u305f\u96c4\u82b1\u306e\u857e\u306e\u5148\u7aef\u3092\u305d\u3063\u3068\u89e6\u308b\u3068\u3001\u307b\u3093\u306e\u6570\u79d2\u3067\u3001\uff11\u8f2a\u306e\u307f\u3042\u308b\u82b1\u88ab\u304c\u80cc\u5074\uff08\u5916\u5074\uff09\u306b\u53cd\u308a\u8fd4\u308a\uff08G\u304b\u3089J\uff1a\uff15\u79d2\u307b\u3069\u3067\u958b\u3044\u305f)\u300120\u672c\u7a0b\u5ea6\u306e\u96c4\u305a\u3044\u304c\u5782\u308c\u4e0b\u304c\u308b\uff08J)\u3002\u96c4\u305a\u3044\u6570\u306f\u82b1\u306b\u3088\u3063\u3066\u5909\u7570\u304c\u3042\u308a\uff08Rutishause\uff52 et al. 1998: \u5185\u8cb4 2017\uff09\u3001\u82b1\u88ab\u304c\uff14\u679a\u306a\u306e\u306b\u3001\u96c4\u305a\u3044\u6570\u304c\uff14\u306e\u500d\u6570\u3067\u306a\u3044\u5834\u5408\u3082\u3042\u308b\u3002\u30a2\u30ab\u30cd\u79d1\u306e\u4ed6\u306e\u7a2e\u3067\u306f\u3001\u96c4\u305a\u3044\u6570\u306f\u7a2e\u5185\u3067\u4e00\u5b9a\u306e\u5834\u5408\u304c\u307b\u3068\u3093\u3069\u3067\u3042\u308b\u304c\u3001\u30e4\u30de\u30c8\u30b0\u30b5\u3067\u3069\u3046\u3057\u3066\u96c4\u305a\u3044\u6570\u304c\u5909\u7570\u3059\u308b\u306e\u304b\u306f\u3088\u304f\u308f\u304b\u3063\u3066\u3044\u306a\u3044\uff08Rutishauser et al. 1998)\u3002 <\/p>\n\n\n\n

When the tip of a mature male flower bud is gently touched, in just a few seconds the single whorl perianth turns abaxially (outward) (G to J: opened in about 5 seconds), and about 20 male stamens hang down (J). The number of stamens varies from flower to flower (Rutishauser et al. 1998: \u5185\u8cb4 2017), and the number of stamens may not be a multiple of 4, even though the flower has 4 perianths. In most other species of the family Rubiaceae, the number of stamens is constant within a species, but it is not well understood why the number of stamens varies in the genus Theligonum<\/em> (Rutishauser et al. 1998).<\/p>\n\n\n\n

\"\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u846f\u7cf8\u3000Filaments<\/a>
\u56f3\uff15\u3000\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u846f\u7cf8\u3000Filaments of Theligonum japonicum<\/em><\/figcaption><\/figure>\n\n\n\n

\u96c4\u305a\u3044\u306e\u82b1\u7cf8(K\u3001L\uff1bL\u306fK\u306e\u9ed2\u8272\u70b9\u7dda\u90e8\u5206\u306e\u62e1\u5927)\u306f\u3001\u9686\u8d77\u304c\u3042\u308a\u7e26\u7e1e\u306e\u3042\u308b\u5947\u5999\u306a\u7d30\u80de\uff08M)\u304c\u3088\u3058\u308c\u308b\u3088\u3046\u306b\u914d\u5217\u3057\u3066\u304a\u308a\u3001\u76f4\u5f8450\u00b5m\u307b\u3069\u3067\u7d30\u3044\u3002\u67d4\u3089\u304b\u304f\u53ef\u52d5\u6027\u304c\u9ad8\u3044\u304c\u3001\u5f37\u5ea6\u304c\u3042\u308a\u5207\u308c\u306b\u304f\u3044\u3002\u305d\u306e\u305f\u3081\u3001\u98a8\u304c\u5439\u304f\u3068\u3001\u846f\u304c\u3086\u3089\u3086\u3089\u63fa\u308c\u3066\u52b9\u7387\u7684\u306b\u82b1\u7c89\u304c\u6563\u5e03\u3055\u308c\u308b\u3002<\/p>\n\n\n\n

The filaments of stamens (K, L; L is an enlargement of the black dotted area of K) is a wiggly array of odd cells (M) with ridges and longitudinal stripes, about 50 \u00b5m in diameter and thin. They are soft and mobile, but strong and hard to break. Thus, when the wind blows, the anthers wobble and effectively disperse pollen.<\/p>\n\n\n\n

\"\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u96cc\u3057\u3079\u3000Female<\/a><\/figure>\n\n\n\n
\"\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u96cc\u3057\u3079\u3000Female<\/a>
\u56f3\uff16\u3000\u30e4\u30de\u30c8\u30b0\u30b5\u306e\u96cc\u3057\u3079\u3000Female flower of Theligonum japonicum<\/em><\/figcaption><\/figure>\n\n\n\n

\u96cc\u82b1\u306f\u751f\u6b96\u30b7\u30e5\u30fc\u30c8\u306e\u8449\u306e\u8449\u814b\u306b\u5f62\u6210\u3055\u308c\u308b\uff08A\u3001B\u3001N-Q\uff09\u3002(Q\u3001R\uff09\u5b50\u623f\u4e0b\u4f4d\u3060\u304c\u3001\u82b1\u88ab\u3068\u96cc\u305a\u3044\u306f\u3001\u4ed6\u306e\u5b50\u623f\u4e0b\u4f4d\u306e\u82b1\u306e\u3088\u3046\u306b\u5b50\u623f\u306e\u5148\u7aef\u3067\u306f\u306a\u304f\u3001\u5b50\u623f\u306e\u5074\u9762\u4e0b\u65b9\u304b\u3089\u4f38\u9577\u3059\u308b\u3002\u82b1\u88ab\u306f\uff11\u8f2a\u3060\u3051\u5f62\u6210\u3055\u308c\u3001\u82b1\u5f01\u3068\u843c\u7247\u306e\u3069\u3061\u3089\u304c\u6d88\u5931\u3057\u305f\u304b\u306f\u308f\u304b\u3063\u3066\u3044\u306a\u3044\u3002\u4ed6\u306e\u30a2\u30ab\u30cd\u79d1\u306e\u7a2e\u3067\u306f\u843c\u7247\u306f\u679c\u6642\u307e\u3067\u5bbf\u5728\u3057\u3001\u82b1\u5f01\u306f\u65e9\u843d\u3059\u308b\u5834\u5408\u304c\u591a\u304f (Robbercht 1988)\u3001Theligonum<\/em>\u306e\u82b1\u88ab\u306f\u3001\u65e9\u843d\u6027\u3067\u3042\u308b\u3053\u3068\u304b\u3089\u3001\u82b1\u5f01\u3067\u306f\u306a\u3044\u304b\u3068\u3044\u3046\u4eee\u8aac\u304c\u63d0\u5531\u3055\u308c\u3066\u3044\u308b\uff08Rutishauser 1998)\u3002\u4e00\u65b9\u3001\u82b1\u5f01\u304c\u9000\u5316\u3057\u3066\u843c\u7247\u306e\u307f\u304c\u6b8b\u3063\u3066\u3044\u308b\u3068\u3044\u3046\u898b\u89e3\uff08\u5185\u8cb4 2017)\u3082\u3042\u308b\u3002\u843c\u7247\u3067\u50cd\u304f\u907a\u4f1d\u5b50\u3001\u82b1\u5f01\u3067\u50cd\u304f\u907a\u4f1d\u5b50\u306e\u3069\u3061\u3089\u304c\u767a\u73fe\u3057\u3066\u3044\u308b\u304b\u306a\u3069\u3001\u3055\u3089\u306a\u308b\u7814\u7a76\u304c\u5fc5\u8981\u3067\u3042\u308b\u3002<\/p>\n\n\n\n

\u82b1\u67f1\uff08Q\u3068R\u306e\u8d64\u8272\u661f\u5370\uff09\u306f\u82b1\u88ab\uff08Q\u306e\u9ed2\u8272\u661f\u5370\u3001R\uff09\u306b\u8986\u308f\u308c\u308b\u90e8\u5206\u306e\u307f\u3067\u3001\u4e73\u982d\u72b6\u7d30\u80de\u3092\u6301\u3064\u67f1\u982d\u304c\u9577\u304f\u4f38\u9577\u3059\u308b\u3002\u5b50\u623f\u306f\u767a\u751f\u521d\u671f\u306f\uff12\u5ba4\u3060\u304c\u3001\uff11\u3064\u306e\u80da\u73e0\u306e\u307f\u304c\u767a\u9054\u3057\uff08S\u3001T)\u3001\u6700\u7d42\u7684\u306b\uff11\u5ba4\u3068\u306a\u308b\uff08Rutishauser et al. 1998)\u3002<\/p>\n\n\n\n

\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u306f\u3001\u4ed6\u306e\u591a\u304f\u306e\u30a2\u30ab\u30cd\u79d1\u306e\u7a2e\u3068\u6bd4\u8f03\u3057\u3066\u3001\uff08\uff11\uff09\u8449\u5e8f\u304c\u5e38\u306b\u5bfe\u751f\u3067\u306f\u306a\u304f\u3001\u6804\u990a\u30b7\u30e5\u30fc\u30c8\u306f\u5bfe\u751f\u3060\u304c\u3001\u82b1\u5e8f\u306f\u4e92\u751f\u3078\u3068\u8ee2\u63db\u3059\u308b\u3053\u3068\u3001\uff08\uff12\uff09\u866b\u5a92\u3067\u306f\u306a\u304f\u98a8\u5a92anemophylly\u3067\u3042\u308b\u3053\u3068\u3001\uff083\uff09\u96c4\u82b1\u3001\u96cc\u82b1\u3068\u3082\u306b\u3001\u82b1\u88ab\u304c\uff12\u8f2a\u3067\u306f\u306a\u304f\uff11\u8f2a\u3067\u3042\u308b\u3053\u3068\u3001\uff08\uff14\uff09\u96c4\u82b1\u304c\u8449\u814b\u3067\u306f\u306a\u304f\u3001\u8449\u814b\u306e\u53cd\u5bfe\u5074\u304b\u3089\u751f\u3058\u308b\u3053\u3068\u3001\uff08\uff15\uff09\u96c4\u82b1\u306e\u96c4\u305a\u3044\u304c\uff11\u8f2a\u306e\u82b1\u88ab\u3068\u540c\u6570\u3067\u306f\u306a\u304f\u30012\u304b\u308919\u672c\u3082\u3042\u308b\u3053\u3068\u3001\uff08\uff15\uff09\u82b1\u67f1\u304c\u307b\u3068\u3093\u3069\u7121\u304f\u3001\u5186\u67f1\u72b6\u306b\u4f38\u3073\u305f\u67f1\u982d\u304c\u5b50\u623f\u57fa\u90e8\u304b\u3089\u5074\u751f\u3059\u308b\u3053\u3068\u3001\uff08\uff16\uff09\u5b50\u623f\u304c\uff12\u5ba4\u304c\u7652\u5408\u3057\u3066\uff11\u5ba4\u3068\u306a\u308a\uff11\u80da\u73e0\u306e\u307f\u5f62\u6210\u3055\u308c\u308b\u3053\u3068\u306a\u3069\u306e\u70b9\u304c\u7570\u306a\u3063\u3066\u3044\u308b\uff08Rutishauser et al. 1998)\u3002\u305f\u3060\u3001\u3069\u306e\u5f62\u8cea\u3082\u30a2\u30ab\u30cd\u79d1\u306e\u4e2d\u3067\u898b\u3089\u308c\u308b\u5f62\u8cea\u3067\u3042\u308a\u3001\u30a2\u30ab\u30cd\u79d1\u306b\u542b\u3081\u308b\u306e\u304c\u59a5\u5f53\u3067\u3042\u308b\u3068\u63d0\u5531\u3055\u308c\u305f\u304c\uff08Wunderlich 1971)\u3001\u7570\u8ad6\u3082\u3042\u3063\u305f\uff08Praglowski 1973, Nowicke and Skvarla 1979)\u3002\u3057\u304b\u3057\u3001\u8a73\u7d30\u306a\u5f62\u614b\u6bd4\u8f03\uff08Rutishauser et al. 1998)\u3001\u5206\u5b50\u7cfb\u7d71\u89e3\u6790\uff08Bremer 2009)\u304b\u3089\u3001\u30a2\u30ab\u30cd\u79d1\u306e\u30a2\u30ab\u30cd\u9023Rubieae\u3001\u30b1\u30ed\u30ae\u30a2\u5c5eKelloggia<\/em>\u3001\u30d7\u30c8\u30ea\u30a2\u9023Putorieae\u3068\u5358\u7cfb\u7d71\u7fa4\u3067\u3042\u308b\u3053\u3068\u304c\u308f\u304b\u3063\u305f\u3002\u30cf\u30b7\u30ab\u30b0\u30b5\u306b\u8449\u306e\u5916\u89b3\u304c\u4f3c\u308b\u304c\u3001\u82b1\u306f\u3082\u3068\u3088\u308a\u3001\u6258\u8449\u5f62\u614b\u3001\u8449\u306e\u6bdb\u306e\u751f\u3048\u65b9\u304c\u7570\u306a\u3063\u3066\u304a\u308a\u3001\u5206\u5b50\u7cfb\u7d71\u89e3\u6790\u304b\u3089\u3082\u59c9\u59b9\u95a2\u4fc2\u3067\u306f\u306a\u3044\uff08Bremer 2009, Neupane et al. 2015)\u3002<\/p>\n\n\n\n

Female flowers are formed in the leaf axils of the reproductive shoot (A, B, N-Q). (Q, R) The ovary is inferior, but the perianths and pistil are formed not at the tip of ovary but the lateral side.<\/p>\n\n\n\n

The perianth is transparent, membranous, and trace-like. The petals are thinner than the sepals in other Rubiaceous species, leading to the hypothesis that the sepals are degenerated (Rutishauser et al. 1998) or that the petals are degenerated and only the sepals remain (\u5185\u8cb4 2017), but which is valid is not known. Further research is needed to determine which genes are expressed, those working in the sepals or those working in the petals.<\/p>\n\n\n\n

The style is the only part covered by the petiole, with a long elongated stigma with papillose cells. The ovary is biloculate in early development, but only one ovule develops (S, T) and eventually becomes unilocular (Rutishauser et al. 1998). Compared to many other species in the family Rubiaceae, the genus Theligonum<\/em> has the following special characteristics: (1) the leaves are not always opposite, the vegetative shoot is opposite, but the inflorescence converts to alternate, (2) it is wind pollinated (anemophily) rather than insect pollinated, (3) both male and female flowers have a single perianth whorl, (4) the male flowers are produced on the opposite side of the leaf, and no subtending leaf, (5) the number of stamens are not equal to that of perianths (four perianths), but 2 to 19, and (6) The ovary is bilocular but changes to actually unilocular with a single ovule. (Rutishauser et al. 1998). However, all of the traits as evolutionary trends are found in other Rubiaceae (Rutishauser et al. 1998), and it was proposed that they should be included in the family (Wunderlich 1971), but there were other hypotheses (Praglowski 1973, Nowicke and Skvarla 1979). However, detailed morphological comparisons (Rutishauser et al. 1998) and molecular phylogenetic analysis (Bremer 2009) have shown that Theligonum<\/em> forms a clade with Rubieae<\/em>, Kelloggia<\/em>, and Putorieae<\/em>. Although the leaves are similar in appearance to those of Neanotis hirsuta <\/em>in the same family, morphology of the flowers, leaf and leaf hair are different, and molecular phylogenetic analysis indicates that they are not sister taxa (Bremer 2009, Neupane et al. 2015).<\/p>\n\n\n\n

\"\u30a2\u30ab\u30cd\u79d1\u3067\u98a8\u5a92\u82b1\u306eCoprosma<\/a>
\u56f3\uff17\u3000\u30a2\u30ab\u30cd\u79d1\u3067\u98a8\u5a92\u82b1\u306eCoprosma repens\u3000<\/em>Wind pollinated flowers of Coprosma repens<\/em> in the Rubiaceae<\/figcaption><\/figure>\n\n\n\n

\u30a2\u30ab\u30cd\u79d1\u3067\u306f\u3001\u30e4\u30de\u30c8\u30b0\u30b5\u9023\u306e\u7a2e\u4ee5\u5916\u306b\u3001Anthospermeae<\/em>\u9023\u3067\u98a8\u5a92\u82b1\u304c\u77e5\u3089\u308c\u3066\u3044\u308b\u3002Anthospermeae<\/em>\u306b\u5c5e\u3059\u308b\u7a2e\u306f\u3001\u4e01\u5b57\u7740\u846f\u3067\u3001\u846f\u7cf8\u304c\u9577\u304f\u3001\u5782\u308c\u4e0b\u304c\u3063\u305f\u96c4\u305a\u3044\u3092\u5f62\u6210\u3057\uff08Rutishauser et al. 1998)\u3001\u98a8\u5a92\u82b1\u3067\u3042\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b\uff08Puff 1982, Thureborn et al. 2019)\u3002\u30a2\u30ab\u30cd\u79d1\u3067\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u4ee5\u5916\u306f\u866b\u5a92\u3067\u3042\u308b\u3068\u8003\u3048\u3089\u308c\u3066\u3044\u308b\u3053\u3068\u304b\u3089\uff08Puff 1982, Thureborn et al. 2019)\u3001Anthospermeae<\/em>\u306e\u5171\u901a\u7956\u5148\u3067\u866b\u5a92\u304b\u3089\u98a8\u5a92\u3078\u306e\u9032\u5316\u304c\u304a\u3053\u3063\u305f\u3068\u63a8\u5b9a\u3067\u304d\u308b\u3002<\/p>\n\n\n\n

Anthospermeae<\/em>\u306e\u7a2e\u3068\u6bd4\u8f03\u3057\u3066\u3001\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u306e\u7a2e\u306f\u3001\u98a8\u5a92\u306b\u5bc4\u4e0e\u3059\u308b\u300c\u591a\u6570\u306e\u300d\u5f62\u8cea\u304c\u9032\u5316\u3057\u3066\u3044\u308b\u70b9\u3067\u7279\u7570\u3067\u3042\u308b\u3002\u591a\u6570\u306e\u96c4\u305a\u3044\u3001\u7d30\u9577\u304f\u3066\u4e08\u592b\u306a\u82b1\u7cf8\u3001\u7d30\u9577\u304f\u958b\u82b1\u5f8c\u3059\u3050\u306b\u88c2\u958b\u3059\u308b\u846f\u3001\u53cd\u308a\u8fd4\u3063\u3066\u96c4\u305a\u3044\u3092\u9732\u51fa\u3055\u305b\u308b\u82b1\u5f01\u3001\u82b1\u67f1\u304c\u77ed\u304f\u67f1\u982d\u304c\u4f38\u9577\u3057\u305f\u96cc\u305a\u3044\u3001\u9000\u7e2e\u3057\u305f\u96cc\u3057\u3079\u306e\u82b1\u88ab\u3001\u96c4\u305a\u3044\u304c\u5f62\u6210\u3055\u308c\u308b\u7bc0\u306b\u306f\u96c4\u305a\u3044\u5074\u306b\u8449\u304c\u5f62\u6210\u3055\u308c\u305a\u3001\u96c4\u305a\u3044\u306e\u82b1\u7c89\u304c\u6563\u5e03\u3055\u308c\u3084\u3059\u304f\u306a\u3063\u3066\u3044\u308b\u751f\u6b96\u30b7\u30e5\u30fc\u30c8\u306a\u3069\u3067\u3042\u308b\u3002\u8907\u5408\u7684\u306a\u5f62\u8cea\u306e\u9032\u5316\u306b\u306f\u3001\u8d85\u907a\u4f1d\u5b50\u5f62\u6210\u304c\u95a2\u308f\u3063\u3066\u3044\u305f\u53ef\u80fd\u6027\u304c\u8003\u3048\u3089\u308c\u3066\u3044\u308b\uff08\u7dcf\u8aac\u3068\u3057\u3066\u3001\u85e4\u539f 2023\uff09\u3002\u30e4\u30de\u30c8\u30b0\u30b5\u5c5e\u306e\u98a8\u5a92\u306b\u95a2\u308f\u308b\u8907\u6570\u306e\u5f62\u8cea\u304c\u3069\u306e\u3088\u3046\u306a\u907a\u4f1d\u5b50\u306b\u3088\u3063\u3066\u5236\u5fa1\u3055\u308c\u3066\u3044\u308b\u304b\u3001\u305d\u308c\u3089\u306e\u907a\u4f1d\u5b50\u304c\u30b2\u30ce\u30e0\u4e0a\u3067\u3069\u306e\u3088\u3046\u306b\u914d\u5217\u3057\u3066\u3044\u308b\u304b\u3092\u7814\u7a76\u3059\u308b\u3053\u3068\u3067\u3001\u8907\u6570\u306e\u9069\u5fdc\u5f62\u8cea\u306e\u9032\u5316\u306b\u3064\u3044\u3066\u65b0\u3057\u3044\u8996\u70b9\u304b\u3089\u306e\u7814\u7a76\u304c\u9032\u5c55\u3059\u308b\u53ef\u80fd\u6027\u304c\u3042\u308b\u3002\u307e\u305f\u3001\u305d\u308c\u305e\u308c\u306e\u5f62\u8cea\u306f\u30a2\u30ab\u30cd\u79d1\u306e\u4ed6\u306e\u9023\u3067\u3082\u898b\u3089\u308c\u308b\u5f62\u8cea\u3067\u3042\u308a\u3001\u30a2\u30ab\u30cd\u79d1\u306e\u5171\u901a\u7956\u5148\u304c\u6301\u3063\u3066\u3044\u305f\u907a\u4f1d\u5b50\u7cfb\u304c\u5916\u9069\u5fdc\u3068\u3057\u3066\u9032\u5316\u3057\u305f\u53ef\u80fd\u6027\u3082\u3042\u308b\u3002<\/p>\n\n\n\n

In addition to the species of Theligonum, wind-pollinated flowers are known in the tribe Anthospermeae of the family Rubiaceae; species belonging to the Anthospermeae have dorsifix anthers, long filaments, and pendulous stamens (Rutishauser et al. 1998) and are considered to be wind-pollinated (Rutishauser et al. Puff 1982, Thureborn et al. 2019). Since all members of the Rubiaceae are considered to be insect-pollinated (Puff 1982, Thureborn et al. 2019), it can be inferred that wind-pollinated flowers evolved at the common ancestor of the Anthospermeae from ancestral insect-pollinated flowers.<\/p>\n\n\n\n

Compared to Anthospermeae species, Theligonum<\/em> species are unique in that they have evolved multiple traits that contribute to wind pollination. These include numerous stamens, long and strong filaments, long and slender anthers that split open soon after flowering, tepals that are recurved to expose the stamens, pistils with the short style and the elongated stigma, reducted pistil perianths, and reproductive shoots in which the nodes where the male flowers are formed have no leaves on the male flower side, facilitating the dispersal of pollen. It is hypothesized that supergene formation may have been involved in the evolution of complex traits (for a review, \u85e4\u539f 2023). Studying what genes control the multiple traits involved in wind pollination in Theligonum<\/em>, and how these genes are arranged in the genome, may advance new perspectives on the evolution of multiple adaptive traits. Each of these traits is also a trait found in other series of the Rubiaceae, and it is possible that the genetic system possessed by the common ancestor of the Rubiaceae evolved as exaptation.<\/p>\n\n\n\n

References:<\/strong><\/p>\n\n\n\n

Antonelli, A. et al.<\/strong> (2021). Settling a family feud: a high-level phylogenomic framework for the Gentianales based on 353 nuclear genes and partial plastomes. Am J Bot 108<\/strong>: 1143\u20131165.<\/p>\n\n\n\n

Bordbar, F., Mirtadzadini, M., and Razafimandimbison, S.G.<\/strong> (2022). Phylogenetic re-assessment of the delimitation of Plocama<\/em> and its species relationships and limits (Rubiaceae, Putorieae<\/em>): resurrection of the monospecific genus Aitchisonia<\/em> and a description of trib. nov. Aitchisonieae<\/em>. Plant Systematics and Evolution 308<\/strong>: 1\u201311.<\/p>\n\n\n\n

Bremer, B. <\/strong>2009. A review of molecular phylogenetic studies of Rubiaceae. Ann. Missouri Bot. Gard. 96: 4-26.<\/p>\n\n\n\n

Chen, J. and Funston, A.M. <\/strong>(2011) Theligonum<\/em> 346-347, Wu, Z. Y., P. H. Raven & D. Y. Hong, eds. 2011. Flora of China. Vol. 19 (Cucurbitaceae through Valerianaceae, with Annonaceae and Berberidaceae). Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis.<\/p>\n\n\n\n

Neupane, S., Dessein, S., Wikstr\u00f6m, N., Lewis, P.O., Long, C., Bremer, B., and Motley, T.J. <\/strong>(2015). The Hedyotis<\/em>–Oldenlandia<\/em> complex (Rubiaceae: Spermacoceae) in Asia and the Pacific: Phylogeny revisited with new generic delimitations. Taxon 64: 299\u2013322.<\/p>\n\n\n\n

Nowicke<\/strong>, J.W. and <\/strong>Skvarla<\/strong>, J.J.<\/strong> (1979) Pollen morphology: the potential influence in higher order systematics. Ann. Missouri Bot. Gard. 66: 633-700.<\/p>\n\n\n\n

Praglowski<\/strong>, J.<\/strong> (1973) The pollen morphology of the Theligonaceae with reference to taxonomy. Pollen & Spores 15: 385-396.<\/p>\n\n\n\n

Puff, C.<\/strong> (1982). The delimitation of the tribe Anthospermeae and its affinities to the Paederieae (Rubiaceae). Botanical Journal of the Linnean Society 84<\/strong>: 355\u2013377.<\/p>\n\n\n\n

Razafimandimbison, S.G., Kainulainen, K., Senterre, B., Morel, C., and Rydin, C.<\/strong> (2020). Phylogenetic affinity of an enigmatic Rubiaceae from the Seychelles revealing a recent biogeographic link with Central Africa: gen. nov. Seychellea<\/em> and trib. nov. Seychelleeae<\/em>. Mol Phylogenet Evol 143<\/strong>: 106685.<\/p>\n\n\n\n

Rutishauser, R., Ronse Decraene, L.P., Smets, E., and Mendoza-Heuer, I. <\/strong>(1998). Theligonum cynocrambe<\/em>: Developmental morphology of a peculiar rubiaceous herb. Plant Syst. Evol. 210: 1\u201324.<\/p>\n\n\n\n

Rydin, C., Wikstr\u00f6m, N., and Bremer, B.<\/strong> (2017). Conflicting results from mitochondrial genomic data challenge current views of Rubiaceae phylogeny. Am J Bot 104<\/strong>: 1522\u20131532<\/p>\n\n\n\n

Thureborn<\/strong>, O., <\/strong>Razafimandimbison<\/strong>, S.G., Wikstr\u00f6m, N., <\/strong>Khodabandeh<\/strong>, A., and Rydin, C.<\/strong> (2019). Phylogeny of anthospermeae of the coffee family inferred using clock and nonclock models. Int J Plant Sci 180<\/strong>: 386\u2013402.<\/p>\n\n\n\n

Wunderlich, R.<\/strong> (1971) Die systematische Stellung von Theligonum.<\/em> Oesterr. Bot. Z. 119: 329-394.<\/p>\n\n\n\n

\u5185\u8cb4\u7ae0\u4e16\uff082017) \u30a2\u30ab\u30cd\u79d1\u3001\u65e5\u672c\u306e\u91ce\u751f\u690d\u7269\u7b2c4\u5dfb\uff08\u5927\u6a4b\u4ed6\u7de8\uff09\u3001\u5e73\u51e1\u793e\u3001pp. 266-293, PL. 206-228. <\/p>\n\n\n\n

\u85e4\u539f\u6674\u5f66\uff082023\uff09\u8d85\u907a\u4f1d\u5b50\u3001\u5149\u6587\u793e\u65b0\u66f8<\/p>\n\n\n\n

https:\/\/www.gbif.org\/species\/2919295: GBIF Secretariat: GBIF Backbone Taxonomy. https:\/\/doi.org\/10.15468\/39omei<\/a> Accessed via https:\/\/www.gbif.org\/species\/<\/a>2919295 on 26th August 2023<\/p>\n","protected":false},"excerpt":{"rendered":"

\u30e4\u30de\u30c8\u30b0\u30b5\u5c5eTheligonum\u306f\u3001\uff14\u7a2e\u304c\u77e5\u3089\u308c\u3066\u3044\u308b\u3002\u65e5\u672c\u3068\u4e2d\u56fd\u6771\u90e8\u306b\u30e4\u30de\u30c8\u30b0 … \u7d9a\u304d\u3092\u8aad\u3080 →<\/span><\/a><\/p>\n","protected":false},"author":1,"featured_media":0,"comment_status":"closed","ping_status":"closed","sticky":false,"template":"","format":"standard","meta":[],"categories":[102,87,167,114,185,182,132,186,170],"tags":[],"_links":{"self":[{"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/posts\/1166"}],"collection":[{"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/posts"}],"about":[{"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/types\/post"}],"author":[{"embeddable":true,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/users\/1"}],"replies":[{"embeddable":true,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=%2Fwp%2Fv2%2Fcomments&post=1166"}],"version-history":[{"count":9,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/posts\/1166\/revisions"}],"predecessor-version":[{"id":1488,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=\/wp\/v2\/posts\/1166\/revisions\/1488"}],"wp:attachment":[{"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=%2Fwp%2Fv2%2Fmedia&parent=1166"}],"wp:term":[{"taxonomy":"category","embeddable":true,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=%2Fwp%2Fv2%2Fcategories&post=1166"},{"taxonomy":"post_tag","embeddable":true,"href":"http:\/\/www.nibb.ac.jp\/plantdic\/blog\/index.php?rest_route=%2Fwp%2Fv2%2Ftags&post=1166"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}