Life-history monographs of Japanese plants. 12: Asarum caulescens Maxim. (Aristolochiaceae)

The life-history characteristics of Asarum caulescens Maxim. (Aristolochiaceae) are described here. This typical summer-green perennial of the Aristolochiaceae is a unique woodland element distributed in central Honshu, that is, south of Fukushima Prefecture to Shikoku and Kyushu, Japan. It grows in the somewhat shady understory of temperate montane deciduous broad-leaved or mixed forests associated with conifers, such as Cryptomeria japonica and Abies firma . One of the unique features of A. caulescens is its rhizome structures in combination with aerial shoots, which play an important role in vegetative propagation. The perennation strategy of A. caulescens is exceedingly complex, producing various combinations of linear or branched rhizomes bearing different numbers of leaves. Rhizomes consist of two or three segments, each 5 mm to 5 cm long as a unit, connected linearly, but also occasionally branched. At the tip of the newly formed rhizome segment, a single leaf or a pair of cordate leaves are formed. Exceedingly complex branching patterns of the rhizome segments were also recognized, forming ramets in late June to early July. Vegetative propagation by ramet formation obviously plays a very important role in the maintenance of local populations.     

乌金草挥发油镇痛作用的研究

采用热板法和醋酸扭体法两种经典的镇痛模型对乌金草挥发油进行了初步的镇痛作用研究。结果表明,乌金草挥发性成分具有显著的镇痛作用,能明显延长小鼠热板痛阈值,有效降低小鼠因醋酸所致扭体反应次数。在给药1mL/kg剂量和2mL/kg剂量时,该挥发油均具有明显的镇痛效果,且有一定的量效关系。     

Floral Biology of Heterotropa tamaensis (Aristolochiaceae) in Japan

Abstract The floral biology of Heterotropa tamaensis (Makino) F. Maekawa was investigated in native populations. The flowers were protogynous, and the stigmas and anthers were distant from one another. These floral characters are significant for cross-pollination, but breeding experiments showed that the flowers are fairly self-compatible and their pollination is obligately dependent upon animal pollen vectors. The most efficient pollinator for H. tamaensis is a fungus gnat, a member of Mycetophylidae. The fungus gnat visited the flowers relatively frequently and often carried many pollen grains on their hairy bodies. Furthermore, their eggs were frequently found within flowers collected from native habitats. The fungus gnats actively moved about within the flowers and often touched the stigmas and anthers with their bodies. Therefore, H. tamaensis seems to be predominantly self-pollinated, though cross-pollination may occur rarely. The presence of fungus gnat eggs in many flowers strengthened Vogel's hypothesis that the flowers of Heterotropa species mimic the basidiomycetes, in which fungus gnats usually oviposit.     

Population change and immature mortality process of Luehdorfia japonica (Lepidoptera: Papilionidae) feeding on an unusual host plant, Asarum caulescens Maxim. (Aristolochiaceae)

A population of the butterfly Luehdorfia japonica dependent on a small stand of an unusual host plant, Asarum caulescens, was found and observed for 5 years. The plant was growing in patchy clumps in a forest and in an adjoining clearing, but the number of leaves in the clearing decreased during the 5 year study period. Butterflies laid eggs mostly in or near the clearing, and the number of eggs steadily decreased year by year, along with the decrease in the number of leaves. The survival rate of the larvae was similar to that of the same species feeding on the usual host plant, Asarum takaoi, in an adjacent habitat. It is considered that A. caulescens is rarely a host of L. japonica because the plant grows in shady conditions and the butterfly prefers a host plant growing in a sunny locations for oviposition.     

A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy

A cladistic analysis of Asarum was conducted to examine relationships among species within the genus and to test the monophyly of several groups of taxa that have often been treated as segregate genera. Thirty-two species were drawn from throughout the range of the genus, representing a broad sample of sections and all segregate genera. The data matrix included 37 characters derived from various aspects of vegetative and floral morphology. A strict consensus of all most parsimonious trees suggests that Asarum s.l. is monophyletic and consists of two main clades: an Asarum clade, which is characterized by connate styles and inferior ovaries, and an Asiasarum-Hexastylis-Heterotropa clade, which is characterized by ridges on the inner perianth surface, dorsal stigmas, and bifid style extensions. The latter is a large and morphologically diverse clade that includes the North American segregate Hexastylis and two Asiatic segregates. Examination of pollination mechanisms in the context of this phylogeny supports the conclusion that herkogamy, and thus obligate insect pollination, is derived from a plesiomorphic condition of autonomous self-pollination. Associated with herkogamy are characters such as glandular trichomes and other ornamentation of the surface of the calyx that probably represent increased specialization to attract insect pollinators. This study also indicates that chromosomal evolution has occurred via aneuploid decrease from an ancestral chromosome number of 2n = 26 to 2n = 24 in Heterotropa. The recognition of two subgenera, subgenus Asarum and subgenus Heterotropa, corresponding to the two clades in the cladistic analysis, is recommended.     

Molecular phylogeny and taxonomic implications of Asarum (Aristolochiaceae) based on ITS and matK sequences

The genus Asarum (Aristolochiaceae) encompasses approximately 120 species from five sections. Taxonomic controversies concerning the genus Asarum and/or its intrageneric classification remain unresolved. In particular, sect. Heterotropa accounts for a large percentage of the genus (80 of 120 species) and is well diverged in the Sino–Japanese Forest subkingdom. Reconstruction of Heterotropa phylogeny and estimation of its divergence times would provide significant insight into the process of species diversity in the Sino–Japanese floristic region. This study encompassed 106 operational taxonomic units (OTUs), and phylogenetic analyses were conducted based on internal transcribed spacer (ITS) and matK sequences. Although the matK sequences provided informative results solely for section Geotaenium, phylogenetic trees based on ITS regions yielded a clear result for several sections. Three sections, Asarum, Geotaenium and Asiasarum, were supported as robust monophyletic groups, whereas Heterotropa had low support. Sect. Hexastylis was revealed to be polyphyletic, suggesting taxonomic reconstruction would be needed. Sect. Heterotropa comprises two clades, which correspond to species distribution ranges: mainland China and the island arc from Taiwan to mainland Japan via the Ryukyu Islands. It is notable that the common ancestry of the latter clade in the eastern Asian islands was highly supported, suggesting that the present species diversity of Heterotropa was initially caused by allopatric range fragmentation in East Asia.     

Concordant clines and significant correlation between floral and pollen characters in Asarum heterotropoides var. heterotropoides (Aristolochiaceae)

To examine geographic differentiation in Asarum heterotropoides var. heterotropoides in Hokkaido Is. in the northern Japan, in which two putative cryptic species have been suspected to exist, extensive and detailed morphological research on 794 individuals from 44 populations throughout Hokkaido Is. was performed. Among the characters examined, the angle between and tip shape of the calyx lobes and the supratecta of the pollen grains were significantly correlated and were found in similar geographic clines. Among them, the pollen showed two discrete states in almost distinct distribution. Multidimensional scaling analysis showed that individuals within each of the two pollen types had different trends in flower characters. Consequently, we assumed a cline from south to north on Hokkaido Is. For the causes of the cline, the two hypotheses were proposed, primary geographic differentiation or extensive introgressive hybridization between two distinct geographical species existing in the past.     

青城细辛的花器官发生

利用扫描电镜观察了青城细辛（Asarum splendens）的花器官发生过程。青城细辛的花器官为轮状结构,向心发生,依次为两轮3基数的花被原基,两轮6基数的雄蕊原基和一轮6基数的心皮原基。两轮花被原基互生,只有外轮（先发生的一轮）花被原基完全发育,而内轮（后发生的一轮）花被原基在发育过程中逐渐退化。两轮雄蕊原基为离心发生：位于内侧的一轮雄蕊原基先发生,每两个原基正对第一轮发生的花被原基,外侧的一轮雄蕊原基后发生,与内轮雄蕊原基互生。心皮与内侧的一轮雄蕊互生。     

The status of Asarum marmoratum (Aristolochiaceae)

Asarum marmoratum Piper, a species with marbled leaves from southern Oregon and northern California, differs fromA. hartwegii. S. Wats. in many respects, and should be recognized as distinct. The species is described, illustrated, and compared to the four other western North American species ofAsarum in a synoptic key.     

马蹄香雌蕊形态及传粉生物学研究

通过对马蹄香(Asarum henryi Oliv.)雌蕊不同发育阶段的整体发育形态观察和组织切片显微结构观察,结果表明,马蹄香雌蕊心皮离生,不分化为柱头、花柱和子房三部分,开花前在心皮腹缝线上部1／2～2／3处分化出数列多细胞表皮毛,构成鸡冠状柱头,表现出原始的雌蕊结构特征。开花习性、传粉试验观察,杂交指数(OCI)及花粉-胚珠比(P／O)测定结果表明,马蹄香植物为兼性自花传粉繁育系统。     

Asarum yentuense sp. nov. (Aristolochiaceae) from Vietnam

A new species, Asarum yentuense N. Tuan & Sasamoto (Aristolochiaceae), is described from Quang Ninh Province, Vietnam. This species belongs to A. subgen. Heterotropa sect. Longistylis and is most similar to Asarum petelotii, A. glabrum, A. reticulatum and A. magnificum var. dinghuense, but is differentiated by morphology, including the length of the calyx tube and lobes, and phenology.     

Chalcone and flavonol glycosides from Asarum canadense (Aristolochiaceae)

Two chalcone glycosides were isolated, together with seven known flavonol glycosides, from the leaves of Asarum canadense. The structures of the chalcone glycosides were established as chalcononaringenin 2',4'-di-O-glucoside and chalcononaringenin 2'-O-glucoside-4'-O-gentiobioside by chemical, UV, FAB MS, 1H and 13C NMR evidence.     

A New Species of Asarum from Hong Kong

One new species of the genus Asarum (Aristolochiaceae) is described fr-om Hongkong, i.e. Asarum hongkongense S. M. Hwang et T. P. Wong Siu.     

Chalcones and other flavonoids from Asarum sensu lato (Aristolochiaceae)

Seventy-five taxa belonging to the genus Asarum sensu lato were studied for their composition of flavonoids. Three chalcones and an aurone were found as major components. The chalcones were identified as chalcononaringenin 2′,4′-di-O-glucoside, 4,2′,4′-tri-O-glucoside, 4-O-glucoside, and the aurone as aureisidin 4,6-di-O-glucoside. The glycoside, 2′,4′-di-O-glucoside was detected in all taxa examined, and is a chemotaxonomical feature of Asarum sensu lato. 4,2′,4′-Tri-O-glucoside was found from the taxa classified into the genera Asiasarum, Geotaenium and Heterotropa by Maekawa's system. On the other hand, the glycoside was not detected from three Asarum sensu stricto species, A. caudigerum, A. caulescens and A. leptophyllum. In contrast, aurone, aureusidin 4,6-di-O-glucoside occurred in two Asarum s.s., A. caulescens and A. leptophyllum. Thus, the Asarum s.s. and other Maekawa's genera, Asiasarum, Geotaenium and Heterotropa could distinguish by the presence or absence of some anthochlor pigments. Other flavonoids were isolated from the selected 18 Asarum species. They were characterized as some flavonol 3- or 3,7-O-glycosides based on kaempferol, quercetin and isorhamnetin, flavone, apigenin 6,8-di-C-glycoside, flavanone, naringenin 5,7-di-O-glucoside, and xanthone, mangiferin.     

A Preliminary Study of the Pollination Ecology of Pedicularis (Scrophulariaceae) in Japan

Abstract An introductory study of the pollination ecology of six Japanese species of Pedicularis conducted in 1982 and 1983 indicated a close parallel between characteristics of floral mechanisms in Japan and those previously investigated in North America. Floral color, floral form, total dissolved solids in floral nectar, and polinator behavior on flowers of Japanese species are similar to those of North American species. Blooming phenology of Japanese species also corresponds to the North American sequence of long-tongued, nectar-foraging queen bumblebees (Bombus Latr.) pollinating nectariferous , earlier-blooming species nototribically and short-tongued, pollen-foraging workers pollinating nectarless, rostrate flowers of later-blooming ones sternotribically. Electrofocus analysis of peroxidase systems in two Japanese species indicated considerable intrapopulational heterozygosity. Nectar component analysis of two transboreal species indicated intraspecific differences in kinds of sugars between widely disjunct Japanese and North American populations. A tentative conclusion is that the Japanese and North American Pedicularis floras have evolved in a parallel manner from Himalayan migrants, while their transboreal elements are in process of vicariant diversification.     

新西兰岩石百合中两种共存的繁育策略：延迟自动自交和欺骗传粉吸引

延迟自动自交和欺骗传粉吸引在被子植物多数类群中有相关报道,但是至今没有发现两种繁育策略在同一物种中共存现象。本研究通过对照试验检测新西兰岩石百合雄蕊附属物和花冠闭合运动是否分别具有欺骗吸引和延迟自交功能。研究结果表明,新西兰岩石百合黄色的雄蕊附属物拟态花粉（或花药）,约93%的昆虫访花行为源自黄色花药附属物的欺骗吸引,雄蕊附属物的报酬拟态功能有效提高昆虫拜访频率并促进异交。同时,研究发现新西兰岩石百合花期结束时花瓣闭合促使雌雄隔离距离的缩小,花瓣自然闭合的花朵平均结籽数（20.62）显著高于闭合前去雄处理花朵（11.79）。我们的结果表明延迟自动自交与欺骗传粉吸引两种繁育策略在新西兰岩石百合中共存。     

13: Lycoris sanguinea Maxim. (Amaryllidaceae)

The life-history characteristics of Lycoris sanguinea Maxim., a bulbous perennial of the Amaryllidaceae, are described here. Its geographical range is in the warm-temperate to cool-temperate zone of Honshu, Shikoku and Kyushu, Japan, extending further to the Korean Peninsula and China. Its growth form and seasonality are very unique, that is, the productive and reproductive phases are separate, which has been observed in other Lycoris species as well. In late March to mid-April for approximately 2 months, only radical leaves appear above ground; these leaves wither and disappear in late April to early May when the canopy of the deciduous trees casts heavy shade on the ground layer; then, in mid- to late-July when the forest floor becomes even darker, only scapes appear above ground, with two to six orange-red flowers borne on the umbels, occasionally forming large clumps here and there on the forest floor. Lycoris sanguinea is a typical insect-pollinated polycarpic perennial that is pollinated by small bees, such as Amegilla florae , and small butterflies, such as Thymelicus leoninus leoninus and Thymelicus sylvaticus sylvaticus . The seed output per plant is not very high, and 6–20 black spherical seeds, each approximately 6 mm in size, are produced.     

Pollen Morphology of Asarum in China

The present paper describes the pollen morphology of 27 species, 4 varieties and 1 form of Asarum from China. The pollen grains were all examined under light microscope and scanning electron microscope. The apertures of pollen grains of Asarum are variable and so distinct as to allow the identification of individual subgenera and species. The exine ornamentation is compound and may be divided into two types: reticulate or cerebelloid under verrucae. The pollen grains of Subgen. Asarum and Subgen. Heterotropa are distinguishable, which supports the taxonomical subdivision based on the gross morphology. The differences in the pollen morphology between A. caulescens Maxim. and A. sieboldii Miq. in China and in Japan are discussed and some comparisons in the pollenmorphology between several species similar in the gross morphology are also made.