Epicuticular wax morphology and the taxonomy ofRosa (sectionCaninae, subsectionRubiginosae)

Classification within the genusRosa is problematic due to ± continuous variation of morphological characters. The dogroses, (Rosa, sectionCaninae) are classified only by their unique meiotic system not by directly visible characters. The intrasectional structure remains insufficient, as the characters (glands, hairs and prickles) are known to exhibit great morphological divergence affected by both genetic plasticity and environmental influences. Taxonomy of dogroses follows a highly artificial system which allows us to identify morphospecies, which are not necessarily evolutionary species. However, SEM-studies in theCaninae revealed that all taxa of the subsectionRubiginosae are characterised by a granule type of epicuticular waxes, whereas members from the other subsections (with three exceptions) form triangular rodlets. Comparison with specimens grown under the same conditions confirmed these findings on natural populations and additionally revealed that following artificial hybridism, offspring develop the maternal type of wax structure due to the matroclinal inheritance in this section.     

Inheritance of growth form and mechanical characters in reciprocal polyploid hybrids of Rosa section Caninae—implications for the ecological niche differentiation and radiation process of hybrid offspring

Within Rosa section Caninae (dogroses) two different growth types exist, arching (L-type) and erect (D-type). Due to the specific mode of chromosome allocation during meiosis, character inheritance in dogroses is maternally skewed. In this paper we try to assess the mode of inheritance of the ‘growth form’ in reciprocal hybrids of R. canina L. and R. rubiginosa L. to predict possible ecological niches and radiation processes of hybridogenic offspring. A quantitative analysis of mechanical properties indicates that neither an expected parentally skewed nor an intermediate habit was expressed by the hybrids, but a dominant L-type as in R. canina, regardless of the parental relationship (pollen-parent or seed-parent). Our results implicate two possible scenarios to be further tested: (a) R. canina acts as a species with a dominant allele for the growth form character. If so, this would be the first evidence of a Mendelian character inheritance in the dogroses. (b) ‘Growth form’ in the dogroses is not a Mendelian character, but a syndrom which realises as an emergent functional property with underlying phenotypic structural differences in stem and wood anatomy.The first two authors contributed equally to the study.     

Different levels of morphometric variation in three heterogamous dogrose species (Rosa sect.Caninae, Rosaceae)

Difficulties in delimiting well-defined entities in the dogroses (Rosa sect.Caninae) has resulted in very variable taxonomic treatments. The present study was undertaken to provide a background for taxonomy as well as plant breeding. Morphometric diversity was analysed on seedlings obtained from field collections in South Sweden of three species,Rosa dumalis, R. rubiginosa andR. villosa. A canonical variates analysis showed that the three species are relatively distinct whereas two subspecies ofR. dumalis were less well discriminated. Analyses of variance demonstrated that intraspecific variation is pronounced inR. dumalis and, to a lesser extent, inR. villosa.     

Geographically differentiating morphology of genetically similar dogroses: consequences of canina meiosis

Despite their genetic homogeneity, many taxonomic species are described as European dogroses (Rosa L. section Caninae (DC.) Ser.) with consistent morphological variability. Here we report a morphometric study of 27 hip and leaf characters of proximate dogrose populations that include Rosa agrestis, a member of the subsection Rubigineae, in Flanders. In principal components analysis one R. agrestis population in Kanne had intermediate morphology between R. canina and R. corymbifera, on the one hand, and three other R. agrestis populations, on the other, suggesting hybridogenic origin. Half-sib R. agrestis seedlings were grown under controlled conditions and their leaf characters were studied. A tendency toward deviating morphology in seedlings from the Kanne population, analogous to the mother plants in the field, and a correlation for specific leaf characters between mother plants and their corresponding seedlings reinforce a genetic basis for the observed divergence in the Kanne population. The assumed hybridogenic mother plants did not produce fewer seeds per hip than the others of the pure populations. In addition, for all sampled dogroses, which included six species, the diameter of the discus was correlated with the number of seeds per hip, whereas the diameter of the orifice was not correlated with this character. This implies that only the diameter of the orifice and not the diameter of the discus might be decisive for species identification among European dogroses.     

Morphological and RAPD markers show a highly skewed distribution in a pair of reciprocal crosses between hemisexual dogrose species, Rosa sect. Caninae

 The dogroses, Rosa sect. Caninae, are polyploid and characterized by their unique meiosis with an unequal number of chromosomes in the male and female gametes. The pollen cells have 7 chromosomes and the egg cells 21, 28 or 35 depending on the ploidy level of the species. The resulting matroclinal inheritance was studied with both morphological and molecular markers in a pair of reciprocal crosses between R. dumalis and R. rubiginosa (2n=35). A canonical discriminant analysis based on seven morphological characters showed only a minor overlapping between the two progeny groups. In addition, the R. dumalis×R. rubiginosa offspring were more heterogeneous than the offspring from the reciprocal cross in each of the characters analysed. Eleven RAPD markers specific for the R. dumalis parent and 10 RAPD markers specific for the R. rubiginosa parent were scored in the offspring. Each of the offspring exhibited either all, or all-but-one, of the seed parent markers. The average number of pollen donor markers found in the offspring was 3.2 (R. dumalis×R. rubiginosa) and 2.7 (R. rubiginosa×R. dumalis). About half of the pollen donor markers were never transmitted to the progeny. This is, to our knowledge, the first time the highly skewed chromosome distribution in Rosa sect. Caninae has been demonstrated with statistically evaluated morphological data and with molecular markers. Received: 25 June 1998 / Accepted: 28 July 1998     

Evidence of apomixis in hemisexual dogroses, Rosa section Caninae

All members of Rosa section Caninae, dogroses are polyploid and characterized by their unbalanced meiosis, which in most cases leads to a pronounced morphological influence from the maternal parent. In a previous investigation on a pair of reciprocal crosses between two species in this section, Rosa dumalis and R. rubiginosa (2n=35), nine offspring plants (approximately 10%) did not receive any of the 21 RAPD markers present in the respective pollen parent. This was interpreted as a possible occurrence of apomixis. These nine plants have now been subjected to a further study with additional markers. Thirteen new RAPD markers showed the same result as in the previous investigation: none of the nine plants inherited any of the pollen donor markers. The reproducibility of the RAPD markers was checked by mixing DNA samples to obtain a series of artificial hybrids between the two parent plants. Twelve RAPD markers gave the expected result, whereas one marker appeared only 50% of the time. In addition, pollen viability, mean number of seeds per hip, mean seed weight, and mean weight of fruit flesh per hip have been studied on the four progeny groups: R. dumalis×R. rubiginosa plants which received pollen donor markers (PM plants), R. dumalis×R. rubiginosa plants which did not receive any pollen donor markers (NPM plants), R. rubiginosa×R. dumalis PM plants and R. rubiginosa×R. dumalis NPM plants. A canonical discriminant analysis based on these four reproductive characters separated the four progeny groups. There were significant differences between the two PM groups in all investigated characters, and also between the PM and the NPM groups in pollen viability. The result from the RAPD markers together with the differences in pollen viability between the PM and NPM progeny groups is taken as an indication that apomixis occurs within the Caninae section. Received: 13 October 1999 / Revision accepted: 28 January 2000     

Reproduction and Hybridisation in the Genus Rosa,Section Caninae (Ser.) Rehd.

Crossing experiments were carried out in order to investigate the importance of apogamy in relation to autogamy, geitonogamy and xenogamy in the reproductive biology of pentaploid members of the genus Rosa, section Caninae (Ser.) Rehd. About 10,000 seeds from 900 crossing experiments were examined, 4500 of them for both filling of the grain and viability of the embryo. The results were compared with experimental reports and literature dating from 1753 to 1995. Apomictic reproduction was shown to be possible in all species of the section tested, but viable seed yields were only about 5% of those produced by xenogamy. This contradicts the hypothesis of Fagerlind (1940,1944,1945,1950) and Custafsson and Hakansson (1942), but supports the results of Matsson (1912) and Schwertschlager (1915). The number of viable seeds increased in the order apogamy, autogamy, geitonogamy to xenogamy. Reciprocal crossings showed significant differences in the number of seeds produced in the hip. This contradicts the hypothesis on the genomic constitution of the pentaploid Caninae with unbalanced heterogamy, as advocated by Hurst (1927) and Custafsson and Hakansson (1942).     

垂叶榕叶片表面蜡质化学组分及蓟马锉吸胁迫对其含量的影响

为明确垂叶榕(Ficus benjamina)叶片表面蜡质的化学组分和蓟马(Thrips spp.)锉吸胁迫对蜡质含量的影响,采用气相色谱-质谱联用仪(GC-MS)对不同危害等级蓟马锉吸叶片的表面蜡质化学组分及含量进行了检测和比较分析。结果表明,垂叶榕叶片的表面蜡质中含有32种化合物,包含有9种酸、8种醇、5种酯、5种酮和5种烃。不同危害等级叶片的表面蜡质总量、已分离化合物的总离子峰面积、各类化合物的相对含量、各类化合物及其主要组分的离子峰面积均不同。垂叶榕叶片的表面蜡质的化学组分和含量可直接或间接反映叶片受蓟马锉吸危害的程度。这说明叶片表面蜡质的化学组分及含量在垂叶榕与蓟马的互作和协同进化中扮演着重要角色。     

Evolutionary patterns and processes in the genus Rosa (Rosaceae) and their implications for host-parasite co-evolution

The genus Rosa (Rosaceae) is a remarkable genus with respect to systematic biology. Multiple reproductive strategies ranging from apomixis to outcrossing including hybridisation, as well as different modes of character inheritance like matroclinal, paternal or sex unrelated ones. This complexity makes the genus a model in which simple concepts of radiation, speciation and taxonomy come to their limits. Analyses of the evolutionary patterns and processes in Rosa give clear evidence for the stochastic character of this specific evolutionary scenario.     

中国17种蔷薇属植物45S和5S的rDNA分布研究

为了探寻蔷薇属植物亲缘关系及系统发育研究的分子细胞遗传学证据,该研究采用双色FISH(荧光原位杂交)技术,对原产中国7个组的17种蔷薇属植物的45S和5S rDNA进行了定位分析。结果表明:(1)多数蔷薇属植物1组染色体对应1个45S rDNA位点和1个或2个5S rDNA位点,偶尔出现1~2个rDNA位点的丢失,但复伞房蔷薇(Rosa brunonii)的1组染色体对应了2个45S rDNA位点。(2)二倍体的蔷薇属植物至少有1对5S rDNA位点与45S rDNA位点共定位,而四倍体材料的5S rDNA位点与45S rDNA位点没有共定位,但所有四倍体材料均至少有1种rDNA信号纯合,表明它们应为二倍体直接加倍产生的同源四倍体。(3)绝大多数材料45S rDNA位于染色体短臂、5S rDNA位于染色体长臂,但缫丝花(R. roxburghii f. roxburghii)有1个5S rDNA信号位于染色体的短臂上,表明它与蔷薇属其他种的亲缘关系较远。(4)阿克苏地区和伊犁地区的疏花蔷薇的核型不同,且45S和5S rDNA的数量和位置不同,分子细胞遗传学证据也支持阿克苏地区的疏花蔷薇应为疏花蔷薇的新变种。(5)该研究中共有8个二倍体和6个四倍体蔷薇属植物的双色FISH为首次报道。研究认为,无论二倍体还是四倍体蔷薇属植物中出现的异形同源染色体、rDNA信号位置在同源染色体上的差异以及rDNA信号的增加和丢失,可能都与染色体结构变异和染色体重组有关,在分子细胞遗传学水平上证明染色体结构变异和染色体重组在蔷薇属植物演化过程中具有重要的作用。     

桉树及其杂交种叶片形态的遗传变异特征

为了解桉树重要性状在杂交种亲本、子代间的遗传变异规律,以尾叶桉(Eucalyptus urophylla)为母本、粗皮桉(E. pellita)和细叶桉(E. tereticornis)为父本、8个全同胞杂交子代为材料,对亲本和子代的叶片形态性状进行了分析。结果表明,亲缘关系近的尾叶桉和粗皮桉的叶片性状大多无显著差异;但他们与亲缘关系远的细叶桉间的差异显著;尾粗桉与父母本间的性状无显著差异或仅与母本差异显著,属偏父本型;尾细桉与双亲间的多数性状都差异显著,且多偏向于母本。在树种内的家系间或个体间,叶片数量性状差异显著,而质量性状仅叶面革质厚度、叶面朝向差异显著;杂交种家系部分性状有超亲现象,且尾细桉家系超亲频率显著高于尾粗桉。叶长、叶长宽比、形状因子、叶色、叶面革质是区别叶片形态特征的决定性状。     

Novel<Emphasis Type="Italic"> eceriferum</Emphasis> mutants in<Emphasis Type="Italic"> Arabidopsis thaliana</Emphasis>

We conducted a novel non-visual screen for cuticular wax mutants in Arabidopsis thaliana (L.) Heynh. Using gas chromatography we screened over 1,200 ethyl methane sulfonate (EMS)-mutagenized lines for alterations in the major A. thaliana wild-type stem cuticular chemicals. Five lines showed distinct differences from the wild type and were further analyzed by gas chromatography and scanning electron microscopy. The five mutants were mapped to specific chromosome locations and tested for allelism with other wax mutant loci mapping to the same region. Toward this end, the mapping of the cuticular wax (cer) mutants cer10 to cer20 was conducted to allow more efficient allelism tests with newly identified lines. From these five lines, we have identified three mutants defining novel genes that have been designated CER22, CER23, and CER24. Detailed stem and leaf chemistry has allowed us to place these novel mutants in specific steps of the cuticular wax biosynthetic pathway and to make hypotheses about the function of their gene products.Abbreviations EMS Ethyl methane sulfonate - SEM Scanning electron microscopy - SSLP Simple sequence length polymorphism - WT Wild type     

Low genetic and morphological differentiation in the European species complex of Rosa sherardii,R. mollis and R. villosa (Rosa section Caninae subsection Vestitae)

The polyploid species complex of Rosa villosa sensu lato (Rosa section Caninae subsection Vestitae) consists of three morphologically similar polyploid species: R. sherardii, R. mollis and R. villosa. Whereas R. sherardii is distributed in central Europe, R. mollis and R. villosa represent a vicariant species pair occurring in northern Europe and in mountains of central and eastern Europe, respectively. In this study we analysed multiple data sets (morphology, cytology, microsatellites, AFLP and plastid DNA sequences) to re‐evaluate the systematics of these species and to examine whether cytological differences are reflected in the taxonomy and geographical distributions. Furthermore, these data sets were used to evaluate hypotheses explaining the vicariant distribution of R. mollis and R. villosa. None of these data sets revealed a clear‐cut differentiation between the species. Cytological and molecular data argued for a discrete taxonomic position of the predominantly pentaploid R. sherardii, but these data did not support a separation between the mostly tetraploid R. mollis and R. villosa. Population genetics revealed that samples of the latter species were assembled according to ploidy, but not to species affiliation or geographical distribution. Thus, we assume that the cytologically polymorphic original species had a continuous range prior to the last glaciation period and survived on nunataks or in non‐glaciated coastal regions in northern Europe, but that it failed to recover its former range after the retreat of the ice sheets. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 240–256.     

Delineating the effects of a plant trait on interactions among associated insects

Plant traits can affect ecological interactions between plants, herbivores, and predators. Our study tests whether reduced leaf wax in peas alters the interaction between the pea aphid ( Acyrthosiphon pisum), a foliar-foraging predator (a lady beetle, Hippodamia convergens) and a ground-foraging predator (a ground beetle, Poecilus scitulus). We performed a 2×2×2 factorial experiment in which wax level, presence of H. convergens, and presence of P. scitulus were manipulated. Experimental arenas consisted of a cage surrounding three pea plants. One plant in each cage was stocked with 15 pea aphids. In greenhouse and field cage experiments, we assessed the effect of each factor and their interactions on aphid density. As in previous studies, H. convergens foraged for aphids more effectively on reduced wax peas than on normal peas. Other interactions among H. convergens, P. scitulus , and A. pisum were the same on both types of peas. We consider how aphid movement, plant growth, and a high frequency of predation by P. scitulus on H. convergens influenced pea aphid density.     

中国蔷薇属植物物种丰富度分布格局及其与环境因子的关系

物种丰富度的大尺度地理格局及其成因是宏观生态学及生物地理学的中心议题之一。蔷薇属(Rosa L.)植物具有很高的经济价值和生态价值,探讨中国蔷薇属植物的丰富度分布格局及其影响因素可为该属植物资源的保护和合理开发利用乃至其系统进化研究提供重要依据。基于蔷薇属植物在中国的15451条分布数据和11种地理、气候等环境因子进行了物种丰富度分析和相关性分析,研究结果显示:(1)蔷薇属植物在中国分布不均匀。在水平方向上,蔷薇属植物于26.19°—34.29°N带内有较高的物种丰富度,之后随着纬度的增加而降低,且随着经度的增加表现为先增加后减少,于99.10°—108.47°E间存在明显的峰值;在垂直方向上,蔷薇属植物的物种丰富度随海拔的增加表现为先增加后减少,956.46—3518.60m范围内的丰富度最高。西南横断山区为蔷薇属物种分布的中心地区,新疆北部及东北长白山周边地区为局部聚集区。(2)蔷薇属物种丰富度与各能量、水分和生境异质性因子均呈正相关关系,与气候稳定性因子呈负相关关系。表明中国蔷薇属植物在水分和热量条件好、气候季节性变化小且生境异质性程度高的地方,有着更高的物种丰富度。(3)蔷薇属...     

Floral development and systematic position of <Emphasis Type="Italic">Arillastrum</Emphasis>, <Emphasis Type="Italic">Allosyncarpia</Emphasis>, <Emphasis Type="Italic">Stockwellia</Emphasis> and <Emphasis Type="Italic">Eucalyptopsis</Emphasis> (Myrtaceae)

We have investigated the floral ontogeny of Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis (of the eucalypt group, Myrtaceae) using scanning electron microscopy and light microscopy. Several critical characters for establishing relationships between these genera and to the eucalypts have been determined. The absence of compound petaline primordia in Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis excludes these taxa from the eucalypt clade. Post-anthesis circumscissile abscission of the hypanthium above the ovary in Stockwellia, Eucalyptopsis and Allosyncarpia is evidence that these three taxa form a monophyletic group; undifferentiated perianth parts and elongated fusiform buds are characters that unite Stockwellia and Eucalyptopsis as sister taxa. No floral characters clearly associate Arillastrum with either the eucalypt clade or the clade of Stockwellia, Eucalyptopsis and Allosyncarpia.We gratefully acknowledge Clyde Dunlop and Bob Harwood (Northern Territory Herbarium) for collecting specimens of Allosyncarpia, and Bruce Gray (Atherton) for collecting specimens of Stockwellia. The Australian National Herbarium (CANB) kindly lent herbarium specimens of Eucalyptopsis for examination. This research was supported by a University of Melbourne Research Development Grant to Andrew Drinnan.     

Systematic importance of pollen morphological features of selected species from the genus Rosa (Rosaceae)

The aim of this study is verification of the taxonomic usefulness of the pollen grain features studied, based on pollen morphology of 32 wild species from all 4 subgenera and all 10 sections of the genus Rosa, mainly for delimitation of subgenera, sections, and species. The measurements and observations were carried out with both light microscopy and scanning electron microscopy. Only correctly formed pollen grains (30 per specimen) were measured, and 960 pollen grains were examined in total. They were analyzed for 14 quantitative features of pollen grains and exine sculpturing and the following qualitative ones: outline, shape, and operculum structure. Our study revealed that the diagnostic features of pollen grains studied were: exine sculpture, length of polar axis, and pollen shape (P/E ratio). On the basis of the above characters, five species were isolated and the remaining ones were included in several groups isolated on the basis of exine sculpture types. The following three exine sculpture types occurred in the species studied: granular-verrucate (in R.?stellata), striate-psilate (in R.?multibracteata and R.?multiflora), and striate (the remaining species). R.?banksiae is characterized by small pollen grains, while R.?setigera has strongly elongated pollen with P/E ratio >1.5. Exine sculpture features considered to be diagnostic should be treated as auxiliary because they fail to differentiate individual species, although they can be helpful in distinguishing groups of species of similar exine sculpture. The arrangement of the species examined on a dendrogram only slightly corroborates division of the Rosa genus into subgenera and sections currently adopted in taxonomy (Rehder 1940). An interesting result was reported for the species studied from the Caninae (R.?agrestis, R.?canina, R.?dumalis, R.?jundzillii, and R.?rubiginosa) section which, despite hybrid nature, with the exception of R.?villosa, grouped in the same, most separated group of species.     

Comparative analysis of micromorphological characters in two distantly related Mexican oaks, Quercus conzattii and Q. eduardii (Fagaceae), and their hybrids

Interspecific hybridization occurs with high frequency in the genusQuercus, but few studies have analyzed and compared micromorphological characters in putative parental species and their hybrids.Quercus eduardii andQ. conzattii are two Mexican black oak species that, although distantly related, have formed at least one population of hybrid origin, where individuals with intermediate macromorphology are present. The purpose of this investigation was to analyze the degree of differentiation in micromorphological characters between the two species and to assess the expression of these characters in individuals with intermediate macromorphology. Foliar trichomes, epicuticular waxes, stomata, and pollen grains, were examined using scanning electron microscopy (SEM) in the three types of individuals (Q. conzatti, Q. eduardii, and intermediates). Trichome density was quantified with light microscopy. Types of trichomes present, length of trichome arms, types of epicuticular wax on the leaf surfaces, and the position of stomata with respect to the foliar surface were characters useful to differentiate betweenQ. conzattii andQ. eduardii. Plants with intermediate macromorphology displayed a pattern of micromorphological characters that were identical to one parental species (Q. conzattii), or extreme or novel relative to both species.