Studies on theStellaria longipes complex (Caryophyllaceae): Isozyme variability and the relationship betweenStellaria longipes andS. longifolia

Genetic variation within and the relationship betweenStellaria longipes Goldie andS. longifolia Muhl. were studied. Ten enzyme systems were assessed in eight natural populations ofS. longipes (25 loci) and three ofS. longifolia (20 loci) using starch and polyacrylamide gel electrophoresis. Patterns of population differentiation corresponded to geographic distance. There was no evidence that polyploidS. longipes had greater electrophoretic variability than diploidS. longipes. The isozyme data confirmed extensive population differentiation in these species and, within that context, a relatively close relationship betweenS. longipes andS. longifolia. It was postulated that diploids of these two species might be the progenitors of tetraploidS. longipes.     

Electrophoretic evidence for the origins of tetraploids and hybrids ofSphenomeris chinensis sensu lato (Lindsaeaceae) in Japan

Hypotheses for the origins of tetraploids and hybrids ofSphenomeris chinensis sensu lato in Japan were tested using electrophoretic analysis of isozymes. Comparison of banding patterns supported the hypothesis that plants having abortive spores had resulted following hybridization between diploidS. chinensis var.chinensis andS. biflora and between diploid var.chinensis andS. gracilis. Results from isozymic analyses also revealed thatS. intermedia probably originated through allotetraploid speciation following hybridization between var.chinensis andS. biflora. There were two electrophoretic variants among tetraploids ofS. chinensis. One variant, restricted to the southern Ryukyu Islands, is probably an allotetraploid of hybrid origin between diploidS. chinensis andS. gracilis. It has been described as a new species,S. yaeyamensis Lin, Kato & K. Iwatsuki. The other variant, morphologically corresponding to var.chinensis, is distributed at least throughout southern Japan and also in northern Thailand. Its origin is uncertain, but one of the parents is probably diploidS. chinensis. Sphenomeris intermedia andS. yaeyamensis occur inland, whereas the parentalS. biflora andS. gracilis are halophilous and rheophllous, respectively.     

DNA contents,giemsa banding,and systematics inScilla bifolia,S. drunensis,andS. vindobonensis (Liliaceae)

DNA contents have been determined cytophotometrically in the three Central European, relatedScilla speciesS. bifolia (2n = 18, 2 x, 1 C = 6.2 pg),S. drunensis (2n = 36, 4 x, 1 C = 12.8 pg), andS. vindobonensis (2n = 18, 2 x, 1 C = 9.4 pg). The tetraploid speciesS. drunensis contains twice as much DNA as the diploidS. bifolia. However, the diploid speciesS. vindobonensis differs in DNA content fromS. bifolia by a factor of about 1.5. This difference is largely due to euchromatic DNA, although the higher DNA content inS. vindobonensis is combined with higher heterochromatin content. The data indicate thatS. bifolia andS. drunensis on the one hand, andS. vindobonensis on the other hand are phyletically well separated. Previous taxonomic conclusions from morphology as well as C-banding are thus corroborated.Evolution ofScilla and Related Genera, V.     

Distinction in morphology and esterase isozyme betweenEupatorium glehni (=E. chinense subsp.sachalinense) andE. chinense var.oppositifolium (compositae)

The taxonomic status ofEupatorium chinese var.oppositofolium andE. glehni (=E. chinense subsp.sachalinense) inE. chinense complex semsu Kitamura has long been controversial. In this paper, the degree of divergence between diploids of these two taxa was examined by means of morphological studies including principal component analysis, the electrophoretic analysis of esterase isozyme variation and observations on habitats. The data obtained through the examinations indicate these two taxa are diverged enough to be recognized as distinct biological species. Since the polyploidE. chinense var.oppositifolium is more or less intermediate in morphology between the two diploid taxa, it is considered to have masked the distinction between the two diploid taxa. Also, electrophoretic evidence suggests that polyploidE. chinense var.oppositifolium is not a hybrid or hybrid derivative withE. glehni as a parental species. Possible origin of polyploidE. chinense varoppositifolium is also discussed.     

Standard karyotypes ofAegilops uniaristata,Ae. mutica,Ae. comosa subspeciescomosa andheldreichii (Poaceae)

C-banding patterns and polymorphisms were analyzed in several accessions of the diploidAegilops speciesAe. uniaristata, Ae. mutica, andAe. comosa subsp.comosa and subsp.heldreichii, and standard karyotypes of these species were established. Variation in C-band size and location was observed between different accessions, but did not prevent chromosome identification. One accession ofAe. uniaristata was homozygous for whole-arm translocations involving chromosomes 1N and 5N. The homoeologous relationships of these chromosomes were established by comparison of chromosome morphologies and C-banding patterns to other diploidAegilops species with known chromosome homoeology. In addition, in situ hybridization analysis with a 5S rDNA probe was used to identify homoeologous groups 1 and 5 chromosomes. The present analysis permitted the assignment of allAe. mutica, comosa subsp.comosa, andAe. comosa subsp.heldreichii chromosomes, and three of the sevenAe. uniaristata chromosomes according to their homoeologous groups. The data presented will be useful analyzing genome differentiation in polyploidAegilops species.     

Chloroplast DNA inheritance in theStellaria longipes complex (Caryophyllaceae)

Inheritance of chloroplast DNA (cpDNA) was examined in F₁ progenies derived from three crosses and three corresponding reciprocal crosses betweenStellaria porsildii andS. longifolia. Chloroplast DNA restriction fragments were analyzed using methods of nonradioactive digoxigenin-11-dUTP labeling and chemiluminescent detection with Lumi-Phos 530. Distinct interspecific restriction fragment polymorphisms were identified and used to demonstrate the mode of cpDNA inheritance. Mode of cpDNA inheritance differed among crosses. Two crosses in whichS. porsildii, SP2920-21, was the maternal parent exhibited three different types of plastids, maternal, paternal and biparental, among the F₁ hybrids, suggesting a biparental cpDNA inheritance and plastid sorting-out inStellaria.     

Stylosanthes sp. aff.S. scabra: a putative diploid progenitor ofStylosanthes scabra (Fabaceae)

Stylosanthes sp. aff.S. scabra is an undescribed taxon showing affinities with the allotetraploid speciesS. scabra, but distinct in a number of attibutes. Several collections show potential as forage for clay soils in northern Australia. Twelve accessions have been analysed using STS (sequence-tagged-sites) as genetic markers, and they all displayed STS phenotypes of typical diploid species. Taking into account their morphological similarities, the STS analysis provides strong evidence thatStylosanthes sp. aff.S. scabra might be a diploid progenitor of the allotetraploidS. scabra. This speculation was supported by cytological examinations. Somatic chromosome numbers of two of these accessions were counted and both were found to be diploid (2n = 20). The level of polymorphism among the 12Stylosanthes sp. aff.S. scabra accessions, estimated using randomly amplified polymorphic DNA (RAPD) as markers, was 7.8%, and the dissimilarity value betweenStylosanthes sp. aff.S. scabra andS. viscosa (the other putative progenitor ofS. scabra) was 89%.     

Chemotaxonomy of theSymphytum officinale agg. (Boraginaceae)

In a chemotaxonomic study of the genusSymphytum pyrrolizidine alkaloids and triterpenes were used as chemotaxonomical markers. A micro-extraction methods was developed for screening compounds of very small pieces of herbarium material. The occurrence of the pyrrolizidine alkaloids symphytine and (acetyl-)lycopsamine is very general forSymphytum taxa. Echimidine is present in someS. officinale L. plants and inS. tanaicense Steven. The triterpene isobauerenol is present inS. officinale, S. bohemicum Schmidt,S. tanaicense and inS. officinale var.lanceolatum Weinm. The chemotaxonomic hypothesis, proposed byGadella and collaborators, based on the presence of the triterpene isobauerenol inS. officinale and its absence inS. asperum Lepech. and the presence of the pyrrolizidine alkaloid echimidine inS. asperum and its absence inS. officinale, can no longer be applied absolutely to theS. officinale species complex. The pyrrolizidine alkaloid and triterpene pattern ofS. officinale (2n = 24) andS. bohemicum (2n = 24) is identical.S. bohemicum is morphologically, cytologically and phytochemically very similar toS. officinale. Furthermore, it readily crosses with the white flowered W. European diploids ofS. officinale. Therefore it seems likely that these two taxa are conspecific.S. tanaicense shows a pyrrolizidine alkaloid and triterpene pattern similar toS. officinale (2n = 40). Also on morphological and cytological grounds they are very similar. It seems highly probable thatS. tanaicense is conspecific withS. officinale (2n = 40) and represents an intraspecific variant only.S. officinale var.lanceolatum contained no pyrrolizidine alkaloids but did contain isobauerenol. This feature points to an origin fromS. officinale.     

Giemsa karyotypes and their evolutionary significance inScilla bifolia,S. drunensis,andS. vindobonensis (Liliaceae)

Quantitatively evaluated C-banded and conventional karyograms are presented for the related speciesScilla bifolia subsp.danubialis Speta,S. drunensis (Speta)Speta, andS. vindobonensis Speta. On the basis of banding patterns and karyotype structureS. bifolia subsp.danubialis (2n = 18, 2×) andS. drunensis (2n = 36, 4×) are quite similar, whileS. vindobonensis (2n = 18, 2×) is entirely different. There is a moderate degree of karyotypic variation withinS. bifolia subsp.danubialis andS. drunensis. However, inS. vindobonensis karyotypes and banding patterns are almost completely stable over a geographical range of about 500 km. The present results confirm the recent taxonomic separation ofS. vindobonensis fromS. bifolia, and suggest a considerable phylogenetic distance between these two diploid species. The results are discussed with reference to the morphological characters of the species and their geographical distribution.Evolution ofScilla and Related Genera, II.Dedicated to Univ.-Prof. Dr.Elisabeth Tschermak-Woess on the occasion of her 60th birthday.     

长柄双花木叶性状异速生长关系随发育阶段和海拔梯度的变化

长柄双花木(Disanthus cercidifolius var. longipes)是一种仅分布于我国东南地区的珍稀濒危植物。为研究该物种叶性状异速生长关系和叶片资源利用策略及其随发育阶段和海拔梯度的变化规律,该文以分布于江西省不同海拔梯度的长柄双花木群落为研究对象,调查分析了群落中不同发育阶段长柄双花木植株的叶片面积、叶片体积以及叶片含水量与叶片干重之间的异速关系。结果表明:不同发育阶段植株之间叶性状异速生长关系有着显著差异。成树叶片面积的增长速度低于或等于叶片干重的增长速度,幼树、幼苗叶片面积的增长速度低于叶片干重的增长速度; 成树叶片体积与叶片干重呈等速增长,幼树、幼苗叶片体积的增长速度高于叶干重的增长速度; 成树叶片含水量的增长速度低于叶干重的增长速度,幼树、幼苗两性状间保持等速增长。海拔梯度对长柄双花木叶性状异速生长关系也有影响,植株叶体积和叶含水量与叶干重的异速生长指数在不同海拔间有显著性差异。在低海拔区域,叶体积与叶干重呈等速增长,叶含水量的增长速度低于叶片干重的增长速度。在高海拔区域,叶体积的生长速度低于叶干重的生长速度,叶含水量和叶片干重呈等速增长。这说明长柄双花木叶片资源投资策略随着发育阶段和海拔梯度的不同发生变化。成树主要将叶生物量投资于光捕获面积和同化结构,幼树和幼苗则主要投资于维管组织的建设。由于海拔升高会引起风力增大、光强增强和土壤理化性质改变,长柄双花木在中低海拔倾向于增大叶体积以抢占资源,在高海拔倾向于加强机械组织和维管组织的建设来抵抗外界因子干扰。     

PATTERNS OF VARIATION IN THE STELLARIA LONGIPES COMPLEX: EFFECTS OF POLYPLOIDY AND NATURAL SELECTION

Patterns of morphological variation were studied in herbarium specimens of Stellaria longipes, an herbaceous perennial and subsequently in a growth chamber experiment using three cytotypes (4x, 6x, 8x) of S. longipes and diploids of its proposed progenitor S. longifolia. Despite extensive phenotypic plasticity in many traits, patterns of variation resulting from ecotypic differentiation within S. longipes could be detected in the field. A distinct form of S. longipes, which is restricted primarily to arctic and alpine tundra locations, shows genetic differentiation for the following traits: few flowers per ramet, a low proportion of flowering ramets, and ovate leaves. The three cytotypes of S. longipes could be distinguished by their mean genotypic value for leaf length and number of flowers per ramet. The extent of phenotypic plasticity in these traits makes it unlikely that the cytotypes could be distinguished in the field. The direction and extent of morphological divergence between S. longifolia and S. longipes suggest an alloploid origin for S. longipes. Variational trends (among-habitat types and cytotypes) in trait means are similar to those reported previously for the pattern of plasticity. This supports the argument that similar forces guide evolution of the mean and pattern of plasticity of a trait.     

The allopolyploid origin of kiwifruit,Actinidia deliciosa (Actinidiaceae)

The genetic origin of kiwifruit (Actinidia deliciosa var.deliciosa) was studied using phylogenetic analysis of DNA sequences derived from the polygalacturonase gene. Results indicate that hexaploid kiwifruit had an allopolyploid origin with the diploidA. chinensis contributing one genome (genome A) and another (as yet unidentified) diploid species contributing a second genome (genome B). The results leave open the question of whether a third, distinct species contributed to the hexaploid kiwifruit genome. A tetraploid race ofA. chinensis is also suggested to be allopolyploid containing genomes A and B.     

Hybridization ofSaxifraga oppositifolia andS. biflora (Saxifragaceae) in a mixed alpine population

Saxifraga oppositifolia, S. biflora, and their hybridS. ×kochii were studied in a mixed population in the Swiss Alps. Qualitative morphological characters did not allow to clearly separate the hybridS. ×kochii from its parental species. All quantitative characters were intermediate inS. ×kochii, but the ranges of these characters overlapped with those ofS. oppositifolia andS. biflora. A hybrid index and canonical discriminant analysis allowed a clear separation of the three groups and strongly confirmed a priori grouping. Phenological observations showed that the three groups were not temporally separated within the investigated mixed population. Reciprocal hand pollinations betweenS. oppositifolia andS. biflora yielded higher seed set withS. oppositifolia than withS. biflora as maternal plant. Germination rate of seeds from these crossing experiments varied in the same way as seed yields, whereas germination rate was intermediate for seeds from naturally pollinatedS. ×kochii. Considering its ecological and reproductive properties,S. ×kochii has good prerequisites for evolving into a distinct species.     

The 5S DNA units of bread wheat (Triticum aestivum)

A collection of 44 cloned 5S DNA units fromTriticum aestivum cv. Chinese Spring were grouped into 12 sequence-types based on sequence similarity and the respective consensus sequences were then produced. The relationship between these 12 consensus sequences (T. aestivum S 1-S 8 andT. aestivum L 1-L 4), together with two clones sequenced byGerlach andDyer, and the 5S DNA consensus sequences from diploidTriticum spp. were then determined by numerical methods. Both phenetic and cladistic analyses were carried out. The following wheat 5S DNA sequences were found to group with respective sequences from diploidTriticum spp.:T. aestivum S 4, S 6 withT. tauschii S;T. aestivum S 3 withT. monococcum S andT. monococcum S-Rus 7;T. aestivum L 1 andT. aestivum L-G&D withT. speltoides L;T. aestivum L 2, L 3 withT. tauschii L;T. aestivum L 4 withT. monococcum L andT. monococcum L-Rus 12. The analyses suggested that 5 out of the 65S Dna loci present in wheat were identified at the sequence level. The locus that could not be identified in this analysis was the5S Dna-B 1 locus. A group ofT. aestivum sequences (T. aestivum S 1, S 7, S 8, S-G&D) were found to be distinct from the other 5S DNA sequences in the data base. The existence of the distinct group of 5S DNA sequences suggests that there is a gap in our current understanding of wheat evolution with respect to the5S Dna loci.     

Culture studies onCaulerpa (caulerpales,chlorophyceae) III. Reproduction,development and morphological variation of laboratory-culturedC. racemosa var.peltata

Reproduction, development and morphological variation of the marine green algaCaulerpa racemosa var.peltata from the southern part of Japan were studied in culture in the laboratory. Anisogamous biflagellate male and female gametes were produced monoeciously and copulated with each other. Settled zygotes became spherical and increased in volume. After five weeks, they formed two germ tubes which extended in opposite directions. Both germ tubes became elongated and branched, resulting in the formation of creeping, filamentous, protonema-like plants. These plants formed primary shoots which differentiated into creeping rhizomes and upright axes. Each upright axis successively formed ramuli and developed into an assimilator. The morphology of assimilators, i.e., shape and arrangement of ramuli, varied with culture coditions. The effects of temperature and light intensity on the formation of assimilators were investigated with 25 combinations of 5 temperatures (20.0–30.0C) and 5 light intensities (0.5–8.0 klux). The morphological plasticity of this alga is identical to that ofCaulerpa racemosa var.laetevirens, which was previously described by the present authors. Thus, apparently, the plasticity of this taxon is correlated with environmental factors. It appears, moreover, thatC. racemosa var.peltata andC. racemosa var.laetevirens are ecophenes (ecads) of a single species.     

Intergeneric hybridization and C-banding patterns inHordelymus (Triticeae,Poaceae)

Crosses ofHordelymus europaeus (2n = 4x = 28) with four genera in theTriticeae were attempted. Adult hybrids were obtained in combinations withHordeum bogdanii (2x),Hordeum depressum (4x), andSecale cereale (2x). The meiotic pairing was very low in the hybrids withH. bogdanii andSecale cereale (0.12 and 0.30 chiasmata/cell, respectively), whereas high pairing (9.90 chiasmata/cell) was found in hybrids withH. depressum due to autosyndetic pairing ofH. depressum chromosomes. The chromosome complement ofHordelymus europaeus comprised 16 metacentrics, 8 submetacentrics, and 4 SAT-chromosomes. The Giemsa C-banding patterns of the chromosomes were characterized by small to minute bands at no preferential positions. It is hypothesized thatHordelymus europaeus may genomically be closest related toTaeniatherum andPsathyrostachys spp.     

A taxonomic study of KoreanSmilacina

Several taxonomic problems of KoreanSmilacina were examined by statistical methods, based on morphological data. Consequently,S. davurica described in Korean flora differs greatly from originalS. davurica, and closely related toS. bicolor morphologically. Therefore, the present authors treated this new taxon as a variety ofS. bicolor, S. bicolor var.flavovirens N. S. Lee et J. Y. Kim var. nov (“Yondusomdae” in Korean). The distribution in South Korea ofS. bicolor var.bicolor is known as only Mt. Chiri, but this species is collected in Mt. Dukyou, also. The bounds of morphological variations ofS. japonica var.japonica, the short plant size and no hairs on the stem, were revealed. The morphological characteristics ofS. japonica var.japonica andS. japonica var.mandshurica is obscure to distinguish. According to the present study,S. japonica var.mandshurica differs fromS. japonica var.japonica in plant size, size of adaxial leaf epidermal cell, ratio of style/ovary length and shape of stigma. The unrecorded taxon in Korean flora,S. robusta, was given a new Korean name as “Kunsomdae.” AlthoughS. japonica var.mandshurica is the variety ofS. japonica var.japonica, it is more closely related toS. robusta than toS. japonica var.japonica.     

Hybridization studies in the genusSilene sectt.Siphonomorpha andAuriculatae (Caryophyllaceae)

Ten species in the genusSilene sectt.Siphonomorpha andAuriculatae were crossed artificially involving 612 crosses to test inter- and infraspecific, intervarietal and intersectional crossability. In sect.Siphonomorpha all interspecific crosses (between diploids) failed due to cross- or seed-incompatibility; however, intervarietal crosses betweenS. gigantea var.gigantea andS. gigantea var.incana produced hybrids. In sect.Auriculatae hybrids were produced betweenS. vallesia andS. boryi, both tetraploids, but crosses between these and the diploid species were unsuccessful. The delimitation and status of the species in both sections was supported by the crossing results.     

自然干旱梯度下的酸枣表型变异

表型变异是植物应对环境变化的一种策略。酸枣植物从中国东部沿海到内陆腹地均有分布,其表型性状的变异可能解释其对自然干旱梯度的适应机制。为验证这一假说,以烟台、石家庄、银川、吐鲁番4个自然干旱梯度生境中生长的酸枣三年生植株的12个表型性状为调查研究对象,采用变异系数和巢式方差分析对酸枣的表型变异进行分析。结果表明:(1)从烟台到银川,叶面积、叶长、叶周长和叶柄长总体呈减小的趋势,而比叶面积呈增大的趋势;(2)随着干旱程度的增强,二次枝的长度、二次枝的基部粗、二次枝的枣吊数、茎比密度和茎水分含量均呈减小趋势,并且种子重和种子短轴长也均呈减小的趋势;(3)对沿干旱梯度分布的4个酸枣种群而言,叶性状的平均变异系数(33.70%)枝性状的平均变异系数(32.41%)种子性状的平均变异系数(9.07%),并且酸枣性状间存在很强的协变。结果表明酸枣的地上部分形态性状沿干旱梯度表现出很强的变异,推测在未来的气候变化下,酸枣将通过这种表型变异的有效组合来适应环境变化。     

Isoenzyme evidence on the systematics ofHordeum sectionMarina (Poaceae)

Morphological differentiation of diploid accessions ofHordeum marinum Huds. s.l. into two varieties, var.marinum and var.fouilladei (Rouy)Nevski is associated with isoenzyme differentiation. The tetraploid form ofH. marinum s.l. exhibited fixed heterozygosity of several heterozymes with one homoeozyme shared with var.fouilladei and the second homoeozyme not found in the two diploids. It also differed from both diploids in the mobility of glucose-6-phosphate dehydrogenase. All three taxa differed in morphs of EST-A. It is concluded that the tetraploid is an allopolyploid with one genome closely related to the diploid var.fouilladei and with the second genome divergent from those of both diploids by genes for unique morphs of eight homoeozymes. On the basis of the isoenzyme data, three phylogenetic sibling species—H. marinum Huds. s.str. (2x),H. geniculatum All. s.str. (= var.fouilladei, 2x), andH. caudatum Jaaska, spec. nova (4x), are proposed within theH. marinum s.l. complex and a key is given.