壳斗科栎属青冈亚属的花粉形态及其系统学意义

对栎属青冈亚属17个种及炭栎花粉形态进行扫描电镜观察。结果表明:青冈亚属和炭栎花粉均为三拟孔沟,球形至长球形,(15～28.0)μm×(15.0～25.6)μm。花粉外壁纹饰可划分为3种:(1)蠕虫状(包括皱波状)、(2)聚合颗粒状、(3)棒状或小刺状。各纹饰在组间镶嵌分布。棒状及小刺状纹饰为栎属花粉较为原始式样,并为青冈亚属与高山栎组及部分巴东组(sect.Cerris)共有,推测其间具有较其它栎类更为接近的亲缘关系,可能为广义栎属中较为基础的类群,而与栎亚属sect.Quercuss.s.,Lobatae和Protobalanus关系较远。花粉形态对广义栎属系统分类价值仅适用于较高分类阶元。     

EXTREME HOST PLANT CONSERVATISM DURING AT LEAST 20 MILLION YEARS OF HOST PLANT PURSUIT BY OAK GALLWASPS

Diversification of insect herbivores is often associated with coevolution between plant toxins and insect countermeasures, resulting in a specificity that restricts host plant shifts. Gall inducers, however, bypass plant toxins and the factors influencing host plant associations in these specialized herbivores remain unclear. We reconstructed the evolution of host plant associations in Western Palaearctic oak gallwasps (Cynipidae: Cynipini), a species-rich lineage of specialist herbivores on oak ( Quercus ). (1) Bayesian analyses of sequence data for three genes revealed extreme host plant conservatism, with inferred shifts between major oak lineages (sections Cerris and Quercus ) closely matching the minimum required to explain observed diversity. It thus appears that the coevolutionary demands of gall induction constrain host plant shifts, both in cases of mutualism (e.g., fig wasps, yucca moths) and parasitism (oak gallwasps). (2) Shifts between oak sections occurred independently in sexual and asexual generations of the gallwasp lifecycle, implying that these can evolve independently. (3) Western Palaearctic gallwasps associated with sections Cerris and Quercus diverged at least 20 million years ago (mya), prior to the arrival of oaks in the Western Palaearctic from Asia 5–7 mya. This implies an Asian origin for Western Palaearctic gallwasps, with independent westwards range expansion by multiple lineages.     

Predispersal reproductive biology of female Osyris quadripartita (Santalaceae), a hemiparasitic dioecious shrub of Mediterranean scrublands

Female Osyris quadripartita plants exhibit uninterrupted reproductive activity throughout the year, due to the long duration of successive stages in the cycle and marked within-crop developmental asynchrony. Cycles corresponding to the flowering seasons of consecutive years overlap in each individual. Flowering takes place in spring, and fruits develop in the dry summer season and ripen at any time of the year. Variation in flowering time explains a negligible proportion of variation in ripening time. The greatest reproductive losses are incurred in the phase extending from closed flowers through unripe fruits, mostly due to ovary abortion. Only 30% of closed flowers eventually reach this latter stage. In contrast, 75% of unripe fruits complete their development, with subsequent dispersal of seeds. The probability of the setting of ripe fruit steadily decreases from early to late season flowers, due to increased ovary abortion rates. Resource limitation in the dry summer season seems responsible for this pattern of selective fruit maturation.     

Phylogeny, biogeography, and processes of molecular differentiation in Quercus subgenus Quercus (Fagaceae).

Quercus is one of the most abundant and economically important genera of woody plants in the Northern Hemisphere. To infer phylogenetic relationships within Quercus subgenus Quercus, chloroplast DNA (cpDNA) restriction sites and nucleotide sequences of the internal transcribed spacers (ITS) and the 5.8S coding region of the nuclear ribosomal DNA repeat were obtained for 44 individuals, including 25 species, intraspecific samples, and three outgroups. Separate parsimony analyses of each data set showed that individual gene trees were congruent and often complementary in supporting clades that generally corresponded to previously recognized taxonomic groups. Only one instance of strongly supported gene tree incongruence was detected and this anomalous pattern was explained best by ancient introgression of cpDNA across sectional boundaries. Simultaneous parsimony analysis of the pruned data sets supported the recognition of the strictly Eurasian section Cerris and resolved a novel hypothesis for the major infrageneric groups (Cerris- (Lobatae- (Protobalanus + Quercus sensu stricto))). The biogeographic hypothesis that all major oak lineages evolved locally at middle latitudes within the general distribution of their fossil ancestors was fully supported. This set of relationships also suggested a New World origin for the widespread white oaks of the Northern Hemisphere (section Quercus s. s.). For both data sets, inter- and intraspecific sampling within section Protobalanus showed little correspondence to morphological species. Greater cladistic structure among the samples was obtained by cpDNA restriction sites and two well-delimited plastomes types comprising a total of 15 distinct haplotypes were resolved. Haplotypes of 2 of the peripheral species in this species complex occupy terminal portions of one of the plastome clades, suggesting a more recent origin relative to those of more widespread species. The phylogeography of the two divergent plastome types suggested a north-south pattern, consistent with a Late Tertiary disjunction in the ancestral distribution of section Protobalanus.     

DEVELOPMENT OF TRISTYLY IN PONTEDERIA CORDATA (PONTEDERIACEAE). I. MATURE FLORAL STRUCTURE AND PATTERNS OF RELATIVE GROWTH OF REPRODUCTIVE ORGANS

Pontederia cordata is a tristylous, self-incompatible emergent aquatic. Measurements of reproductive organs in mature flowers and developing buds and analysis of relative growth rates of the styles, filaments, and floral tube were used to analyze the developmental processes that result in reciprocal positioning of reproductive parts among the style morphs. Flowers are trimerous with two series of three tepals, two series of three stamens, and a tricarpellate ovary with a single ovule. Each flower has stamens of two lengths, but the two lengths correspond to upper vs. lower position in the zygomorphic flowers, not to the two series initiated in the primordium. Morph-dependent variation in stigma height depends on differences in style length, not ovary length. Differences among morphs in anther height result from differences in the position of filament insertion on the floral tube and differences in filament length. Styles of the three morphs develop to different lengths as a result of two distinct processes. Styles of L and M morphs develop at different rates with the L style growing more rapidly than the M style, whereas styles of the S morph have a brief period of rapid growth followed by early inhibition. Stamen growth in the S morph also differs qualitatively from that in the L and M morphs, which are distinguished from each other by quantitative differences in growth rates. Results indicate that the developmental processes that result in the complementary arrangements of organs in different morphs are morph-specific. An attempt is made to integrate the findings of this study with the model for genetic control of tristyly.     

Disruption of endosperm development: an inbreeding effect in almond (Prunus dulcis)

A homozygous self-compatible almond, originated from self-fertilization of a self-compatible genotype and producing a reasonable yield following open pollination, exhibited a very high fruit drop rate when self-pollinated. To investigate whether fruit dropping in this individual is related to an abnormal development of the embryo sac following self-fertilization, histological sections of ovaries from self and cross-pollinated flowers were observed by light microscopy. Additionally, the presence of pollen tubes in the ovary and fruit set were determined for both types of pollination. Despite pollen tubes reached the ovary after both pollinations, differences in embryo sac and endosperm development after fertilization were found. Thus, while for cross-fertilized ovules a pro-embryo and an endosperm with abundant nuclei were generally observed, most self-fertilized ovules remained in a previous developmental stage in which the embryo sac was not elongated and endosperm nuclei were absent. Although 30 days after pollination fruit set was similar for both pollination types, at 60 days it was significantly reduced for self-pollination. These results provide evidence that the high fruit drop in this genotype is the consequence of a disrupted development of the endosperm, what could be an expression of its high level of inbreeding.     

Abnormal development of floral meristem triggers defective morphogenesis of generative system in transgenic tomatoes

Parthenocarpy and fruit malformations are common among independent transgenic tomato lines, expressing genes encoding different pathogenesis-related (PR) protein and antimicrobal peptides. Abnormal phenotype developed independently of the expression and type of target genes, but distinctive features during flower and fruit development were detected in each transgenic line. We analyzed the morphology, anatomy, and cytoembryology of abnormal flowers and fruits from these transgenic tomato lines and compared them with flowers and fruits of wild tomatoes, line YaLF used for transformation, and transgenic plants with normal phenotype. We confirmed that the main cause of abnormal flower and fruit development was the alterations of determinate growth of generative meristem. These alterations triggered different types of anomalous growth, affecting the number of growing ectopic shoots and formation of new flowers. Investigation of the ovule ontogenesis did not show anomalies in embryo sac development, but fertilization did not occur and embryo sac degenerated. Nevertheless, the ovule continued to differentiate due to proliferation of endothelium cells. The latter substituted embryo sac and formed pseudoembryonic tissue. This process imitated embryogenesis and stimulated ovary growth, leading to the development of parthenocarpic fruit. We demonstrated that failed fertilization occurred due to defective male gametophyte formation, which was manifested in blocked division of the nucleus in the microspore and arrest of vegetative and generative cell formation. Maturing pollen grains were overgrown microspores, not competent for fertilization but capable to induce proliferation of endothelium and development of parthenocarpic ovary. Thus, our study provided new data on the structural transformations of reproductive organs during development of parthenocarpic fruits in transgenic tomato.     

The relationship between phenology of pistillate flower organogenesis and mode of heterodichogamy in Juglans regia L. (Juglandaceae)

 We investigated the degree of organogenesis completed at the end of the growing season in pistillate flowers of heterodichogamous Juglans regia, English or Persian walnut. Terminal buds from paired cultivars, one each protandrous and protogynous, chosen to represent early, midseason and late leafing walnuts, were examined by scanning electron microscopy. Results indicate that pistillate floral primordia in protandrous individuals had not progressed beyond involucre initiation during the season prior to bloom. In protogynous individuals, floral differentiation had progressed to the initiation of perianth primordia. These observations are compared with an earlier report on staminate flower differentiation in the same cultivars where a comparable, but opposite, relationship exists. We conclude that the degree of differentiation in both staminate and pistillate flowers that must be completed between the time growth resumes in the spring and anthesis is a developmental determinant of the mode of heterodichogamy in walnut. Received: 15 June 1996 / Revision accepted: 25 October 1996     

Anatomical and developmental study of petrified Quercus (Fagaceae) fruits from the Middle Miocene, Yakima Canyon, Washington, USA

The first reported petrified acorns to show internal anatomical structure are here described from Middle Miocene (～15.6 million years old) chert of the Columbia River Basalt Group in Yakima Canyon, Washington. Quercus hiholensis Borgardt et Pigg sp. nov. is described from anatomical and morphological fruit features, as well as a little recognized anatomical feature, the umbilical complex. Acorns, each comprising a nut and its cupule, are up to 15.3 mm long and 18.8 mm wide with helically arranged, imbricate, tuberculate cupule scales. They show basal aborted ovules, short styles, broad stigmas, and lack grooves in their cotyledons. These characters and the developmental pattern seen in these fossil acorns demonstrate that Q. hiholensis conforms to genus Quercus (Fagaceae), subgenus Quercus, section Quercus (the white oaks). The correspondence of Q. hiholensis to the modern section Quercus reveals that the derived floral and fruit characters that distinguish section Quercus within the genus had evolved by the Middle Miocene.     

Multiple nuclear genes stabilize the phylogenetic backbone of the genus Quercus

Phylogenetic relationships among 108 oak species (genus Quercus L.) were inferred using DNA sequences of six nuclear genes selected from the existing genomic resources of the genus. Previous phylogenetic reconstructions based on traditional molecular markers are inconclusive at the deeper nodes. Overall, weak phylogenetic signals were obtained for each individual gene analysis, but stronger signals were obtained when gene sequences were concatenated. Our data support the recognition of six major intrageneric groups Cyclobalanopsis, Cerris, Ilex, Quercus, Lobatae and Protobalanus. Our analyses provide resolution at deeper nodes but with moderate support and a more robust infrageneric classification within the two major clades, the ‘Old World Oaks’ (Cyclobalanopsis, Cerris, Ilex) and ‘New World Oaks’ (Quercus, Lobatae, Protobalanus). However, depending on outgroup choice, our analysis yielded two alternative placements of the Cyclobalanopsis clade within the genus Quercus. When Castanea Mill. was chosen as outgroup, our data suggested that the genus Quercus comprised two clades corresponding to two subgenera as traditionally recognized by Camus: subgenus Euquercus Hickel and Camus and subgenus Cyclobalanopsis Øersted (Schneider). However, when Notholithocarpus Manos, Cannon and S. Oh was chosen as an outgroup subgenus Cyclobalanopsis clustered with Cerris and Ilex groups to form the Old World clade. To assess the placement of the root, we complemented our dataset with published data of ITS and CRC sequences. Based on the concatenated eight gene sequences, the most likely root position is at the split between the ‘Old World Oaks’ and the ‘New World Oaks’, which is one of the alternative positions suggested by our six gene analysis. Using a dating approach, we inferred an Eocene age for the primary divergences in Quercus and a root age of about 50–55 Ma, which agrees with palaeobotanical evidence. Finally, irrespective of the outgroup choice, our data boost the topology within the New World clade, where (Protobalanus + Quercus) is a sister clade of Lobatae. Inferred divergence ages within this clade and the Cerris–Ilex clade are generally younger than could be expected from the fossil record, indicating that morphological differentiation pre-dates genetic isolation in this clade.     

Reproductive cycle of the blacknose shark Carcharhinus acronotus in the Gulf of Mexico

The reproductive periodicity of blacknose sharks Carcharhinus acronotus in the northern Gulf of Mexico was determined by examining reproductive tissues from specimens collected monthly from 2002 to 2005. Monthly changes in shell gland mass, right ovary mass and ovarian follicle diameter were assessed for 74 mature females. Temporal changes in testes mass, testes width and proportion of mature spermatocysts were examined for 64 mature males. Trends in female reproductive tissues suggested an annual peak in reproductive activity during June and July, while trends in male variables suggested an annual reproductive peak during May and June. Although male and female reproductive activity peaked in different months, a strong synchronicity existed between the proportion of mature spermatocysts and the diameter of the largest ovarian follicle. Based on these results, the mating season of blacknose sharks lasts from mid-May to July in the Gulf of Mexico. Maximum embryo sizes were observed in May, which suggested that partition occurs during late May or early June. Results indicate that blacknose sharks have a clearly defined annual cycle in the Gulf of Mexico. This conclusion is further supported by the complete absence of gravid females without vitellogenic ovarian follicles among all mature females examined.     

海南山薯雌花发育及胚胎发育的研究

通过研究山薯的雌花及胚胎发育,为山薯的胚胎学研究以及杂交育种奠定基础。结果表明:山薯大部分为雌雄异株,海南岛的山薯雌花花期约3个月,为9月初至11月末。子房3室,每室有2个倒生胚珠;胚珠具厚珠心,双珠被。珠孔一端表皮下的孢原细胞逐渐发育为大孢子母细胞。大孢子母细胞减数分裂形成4个呈线形排列的大孢子,其中只有1个可以发育为功能大孢子。成熟的胚囊为7胞8核胚囊,其胚囊发育类型为蓼型。卵细胞的受精属于有丝分裂前型。其胚的发育类型为柳叶菜型,经过二细胞原胚、倒T型原胚、棒状胚、球形胚和梨形胚这5个发育阶段。胚乳的发育为核型。     

中国石龙子繁殖季节血浆性类固醇激素水平的变化

本文研究了中国石龙子繁殖季节中性腺变化与血浆性类固醇激素水平波动间的相互关系。结果表明：雌性中国石龙子卵黄生成需要雌二醇(17β-estradiol,E2)的刺激。体现在卵巢重量于4月底至5月中旬间达峰值,血浆E2水平于3月底至最高值,此后显著回落,并于排卵前再次回升;血浆孕酮(progesterone,P)的生理功能在于输卵管卵的维持、促进胚胎发育和卵壳的形成。从3月中旬至4月底间,血浆P水平在基础值附近波动。5月中旬,随动物的排卵,血浆P水平迅速上升,至该月底达最高值,于产卵后迅速下降;E2随P的上升而下降,两者呈反向变动。雄性血浆睾酮(testosterone,T)水平与生精作用、维持攻击、求偶及交配行为密切相关。血浆T水平于动物出眠后呈现显著的上升趋势,并于4月中旬达峰值。5月中旬后,随睾丸的萎缩,血浆T水平也缓慢下降,至6月底达最低值。     

Changes in competence for somatic embryogenesis in Norway spruce zygotic embryo segments

Embryogenic capacity of Norway spruce zygotic embryo sections was drastically altered by a preinduction transfer to hormone-free medium for 7 or 14 days. An increase in competence for somatic embryogenesis was found with the cotyledons, while the hypocotyl sections completely lost their competence. These changes in competence were not dependent on physical contact between plant sections, and could not be correlated to the developmental stage of each section. The increase of competence in cotyledonary material was due not only to an increase of genotypes initiating somatic embryos, but also to an increase in embryogenicity of cotyledons.     

Self-sterility in Ipomopsis aggregata (Polemoniaceae) is due to prezygotic ovule degeneration

Based on previous studies, extreme (>99%) self-sterility in scarlet gilia (Ipomopsis aggregata) appears to be involved in late-acting ovarian self-incompatibility (OSI). Here, we confirm this suggestion by comparing structural events that follow from cross- vs. self-pollinations of I. aggregata. Growth of cross- and self-pollen tubes in the style at 11 h and growth in the ovary at 24 h was equivalent. Nonetheless, by 24 h, cross-pollen effected a significantly higher percentage of both ovule penetration and fertilization. Ovules in self-pollinated flowers showed pronounced changes, including an absence of embryo sac expansion and reduced starch in the integument, by 11 h post-pollination, well before pollen tube entry into the ovary. In addition, the integumentary tapetum and adjacent 1-3 cell layers exhibited abnormal cell division, pronounced deposition of thick, pectin-rich cell walls, and cellular collapse. Ovules and embryo sacs from cross-pollinated flowers rarely showed such features. Developmental changes in ovules from self-pollinated flowers eventually resulted in integument and embryo sac collapse, a process not observed in ovules of unpollinated flowers. We suggest that OSI involves long-distance signaling between self-pollen or self-pollen tubes and carpel tissue that reduces availability of receptive ovules for fertilization before pollen tubes arrive in the ovary.     

Analysis of developmental preformation in the alpine herb Caltha leptosepala (Ranunculaceae)

Developmental preformation is ubiquitous among alpine and arctic tundra plant species and may cause a delay in plant morphological responses to environmental variation. The duration of preformation and seasonal pattern of development were examined in Caltha leptosepala to identify characteristics of architecture and development that may influence the timing of plant responses to environmental cues, both within a single growing season and between years. All structures in C. leptosepala are preformed: leaves are initiated one or two growing seasons before they mature and flowers are initiated one growing season before maturation. Features of development and architecture in C. leptosepala, however, appear to differ from the determinate growth patterns of other exclusively preforming species, and may allow within-season variability in the seasonal development and maturation of structures. Cohorts of leaves initiated are asynchronous with maturation cohorts, and each year the number of leaf primordia per plant at snowmelt exceeds the number to mature aboveground. Therefore, some flexibility in whether leaves complete a 2-yr or 3-yr developmental trajectory might occur. Plasticity in reproductive phenotype might also occur via the process of floral abortion. Despite developmental characteristics that might facilitate the expression of phenotypic plasticity, only slight variability was observed in the duration of preformation or in the seasonal pattern of initiation and emergence of structures. Growth patterns of C. leptosepala thus appear to be fundamentally constrained, and limitations to annual growth may assure that sufficient preformed primordia remain belowground at the end of each growing season for maturation of a full cohort during the subsequent season.     

一叶萩大孢子发生和雌配子体发育

采用石蜡切片法对一叶萩(Flueggea suffruticosa(Pall.)Baill.)雌花大孢子发生及雌配子体的发育过程进行了观察。结果表明:一叶萩子房上位,中轴胎座,三室,每室2枚倒生胚珠,双珠被,厚珠心,具大戟科常见的珠心喙和珠孔塞。胚囊发育为蓼型,成熟胚囊结构为:卵器位于珠孔端,包括2个助细胞和1个卵细胞;中央是1个二核的中央细胞;合点端为3个反足细胞。本研究为大戟科植物生殖生物学和传粉生物学研究提供了基础资料,同时对大戟科植物系统分类研究有一定意义。