准噶尔无叶豆片断化居群的遗传变异及克隆多样性

准噶尔无叶豆(Eremosparton songoricum)是豆科无叶豆属小半灌木, 既能开花结实进行有性繁殖, 又可以靠根茎进行无性克隆繁殖, 为国家三级保护植物, 在中国仅片断化分布于新疆古尔班通古特沙漠局部区域。本文采用ISSR分子标记对采自古尔班通古特沙漠腹地及边缘的7个准噶尔无叶豆自然居群共148个个体进行了遗传变异和克隆多样性分析。8个引物共扩增出84个位点, 其中77个为多态性位点, 物种水平上的多态位点百分比PPB为91.67％, Nei's基因多样性指数I为0.3192, Shannon信息指数H为0.3540; 居群水平上的多态位点百分比PPB为58.45％, Nei's基因多样性指数I为0.2248, Shannon信息指数H为0.3270。居群间的遗传分化系数G_ST为0.2978。AMOVA分析表明, 有31.88%的遗传变异存在于居群间, 表明居群间存在显著的遗传分化。克隆多样性分析表明, 居群水平上, 居群G的Simpson多样性指数和均匀度指数最高, 分别为0.9400和0.9885; 居群E最低, 分别为0.8457和0.9021。物种水平上, Simpson多样性指数为0.9858, 均匀度指数为0.9673。本研究结果表明, 和其他荒漠植物相比, 准噶尔无叶豆表现出较高的遗传变异水平和克隆多样性, 这主要与该物种兼性生殖的繁育方式及多克隆起源有关; 而居群间产生了显著的遗传分化则主要由于人为干扰引起的生境片断化和居群减小而导致的基因交流障碍所致。遗传变异水平和遗传结构的研究将为分析准噶尔无叶豆致濒原因及进化潜力提供重要资料, 对该物种保护具有指导意义。     

鄂尔多斯高原固沙禾草沙鞭种子休眠和萌发与环境的关系

沙鞭是鄂尔多斯高原流动沙丘上分布的多年生禾草。新采收的种子能够在15～35℃的光照和暗中萌发。在25℃光照下,通过14d的培养,种子的萌发率能够达到80％。在25～35℃温度范围内,种子的萌发在光照下比暗中好。通过低温层积处理表明,沙鞭的种子处于非深度休眠状态。4星期的低温层积处理能够有效地加速和提高种子在20～30℃、光照和暗中萌发。对低温贮藏的需求可能是种子对其生境的适应。另外,划伤颖果的果皮和种皮,以及不同程度地部分移走胚乳也能够不同程度地加速和提高种子的萌发,但幼苗的干重以及根和苗的长度显著地受到了移走胚乳的影响。     

Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.     

Intermediate complex morphophysiological dormancy in seeds of the cold desert sand dune geophyte Eremurus anisopterus (Xanthorrhoeaceae; Liliaceae s.l.)

Background and Aims

Little is known about morphological (MD) or morphophysiological (MPD) dormancy in cold desert species and in particular those in Liliaceae sensu lato, an important floristic element in the cold deserts of Central Asia with underdeveloped embyos. The primary aim of this study was to determine if seeds of the cold desert liliaceous perennial ephemeral Eremurus anisopterus has MD or MPD, and, if it is MPD, then at what level.

Methods

Embryo growth and germination was monitored in seeds subjected to natural and simulated natural temperature regimes and the effects of after-ripening and GA₃ on dormancy break were tested. In addition, the temperature requirements for embryo growth and dormancy break were investigated.

Key Results

At the time of seed dispersal in summer, the embryo length:seed length (E:S) ratio was 0·73, but it increased to 0·87 before germination. Fresh seeds did not germinate during 1 month of incubation in either light or darkness over a range of temperatures. Thus, seeds have MPD, and, after >12 weeks incubation at 5/2 °C, both embryo growth and germination occurred, showing that they have a complex level of MPD. Since both after-ripening and GA₃ increase the germination percentage, seeds have intermediate complex MPD.

Conclusions

Embryos in after-ripened seeds of E. anisopterus can grow at low temperatures in late autumn, but if the soil is dry in autumn then growth is delayed until snowmelt wets the soil in early spring. The ecological advantage of embryo growth phenology is that seeds can germinate at a time (spring) when sand moisture conditions in the desert are suitable for seedling establishment.     

Trade-offs between seed dispersal and dormancy in an amphi-basicarpic cold desert annual

Background and Aims

Several studies have demonstrated trade-offs between depth of seed dormancy and dispersal ability for diaspore-dimorphic species. However, relatively little is known about trade-offs between these two life history traits for a species that produces more than two diaspore morphs. The aim of this study was to investigate the relationship between seed dormancy and dispersal in Ceratocarpus arenarius, an amphi-basicarpic cold desert annual that produces a continuum of dispersal unit morphs.

Methods

A comparison was made of dispersal and dormancy breaking/germination responses of dispersal units from ground level (a), the middle of the plant canopy (c) and the top of the plant canopy (f). Various features of the morphology and mass of dispersal units and fruits (utricles) were measured. The role of bracteoles in diaspore dispersal by wind, settlement onto the soil surface and dormancy/germination was determined by comparing responses of intact dispersal units and fruits. Movement of dispersal units by wind and animals, seed after-ripening, germination phenology and the presence of water-soluble germination inhibitors in bracteoles were tested using standard procedures.

Key Results

Dispersal units a, c and f differed in morphology and mass; in the majority of cases, extremes were exhibited by a and f, with c being intermediate. Overall, relative dispersal ability was f > c > a, whereas relative intensity of dormancy was a > c > f. Bracteoles increased dispersal distance by wind, enhanced settlement of diaspores onto the soil surface and mechanically inhibited germination.

Conclusions

The results provide evidence for a model in which there is a continuous inverse-linear relationship between diaspore dispersal ability and depth of dormancy. Thus, dispersal unit heteromorphism of C. arenarius results in a continuum, from no dispersal ability/high dormancy (dispersal unit a) to high dispersal ability/low dormancy (unit f), which may be a bet-hedging strategy in the cold desert environment.     

Fruit and seed heteromorphism in the cold desert annual ephemeral Diptychocarpus strictus (Brassicaceae) and possible adaptive significance

Background and Aims

Diptychocarpus strictus is an annual ephemeral in the cold desert of northwest China that produces heteromorphic fruits and seeds. The primary aims of this study were to characterize the morphology and anatomy of fruits and seeds of this species and compare the role of fruit and seed hetermorphism in dispersal and germination.

Methods

Shape, size, mass and dispersal of siliques and seeds and the thickness of the mucilage layer on seeds were measured, and the anatomy of siliques and seeds, the role of seed mucilage in water absorption/dehydration, germination and adherence of seeds to soil particles, the role of pericarp of lower siliques in seed dormancy and seed after-ripening and germination phenology were studied using standard procedures.

Key Results

Plants produce dehiscent upper siliques with a thin pericarp containing seeds with large wings and a thick mucilage layer and indehiscent lower siliques with a thick pericarp containing nearly wingless seeds with a thin mucilage layer. The dispersal ability of seeds from the upper siliques was much greater than that of intact lower siliques. Mucilage increased the amount of water absorbed by seeds and decreased the rate of dehydration. Seeds with a thick mucilage layer adhered to soil particles much better than those with a thin mucilage layer or those from which mucilage had been removed. Fresh seeds were physiologically dormant and after-ripened during summer. Non-dormant seeds germinated to high percentages in light and in darkness. Germination of seeds from upper siliques is delayed until spring primarily by drought in summer and autumn, whereas the thick, indehiscent pericarp prevents germination for >1 year of seeds retained in lower siliques.

Conclusions

The life cycle of D. strictus is morphologically and physiologically adapted to the cold desert environment in time and space via a combination of characters associated with fruit and seed heteromorphism.     

Seed development in Ipomoea lacunosa (Convolvulaceae), with particular reference to anatomy of the water gap

BACKGROUND AND AIMS: Disruption of one or both of the bulges (water gap) in the seed coat adjacent to the micropyle is responsible for breaking physical dormancy (PY) in seeds of Ipomoea lacunosa and other taxa of Convolvulaceae. Hitherto, neither ontogeny of these bulges nor onset of PY together with anatomical development and maturation drying of the seed had been studied in this family. The aims of this study were to monitor physiological and anatomical changes that occur during seed development in I. lacunosa, with particular reference to ontogeny of the water gap. METHODS: Developmental anatomy (ontogeny) of seed coat and dry mass, length, moisture content, germinability and onset of seed coat impermeability to water were monitored from pollination to seed maturity. Blocking/drying and dye-tracking experiments were done to identify site of moisture loss during the final stages of seed drying. KEY RESULTS: Physiological maturity of seeds occurred 22 d after pollination (DAP), and 100 % of seeds germinated 24 DAP. Impermeability of the seed coat developed 27-30 DAP, when seed moisture content was 13 %. The hilar fissure was identified as the site of moisture loss during the final stages of seed drying. The entire seed coat developed from the two outermost layers of the integument. A transition zone, i.e. a weak margin where seed coat ruptures during dormancy break, formed between the bulge and hilar ring and seed coat away from the bulge. Sclereid cells in the transition zone were square, whereas they were elongated under the bulge. CONCLUSIONS: Although the bulge and other areas of the seed coat have the same origin, these two cell layers underwent a different series of periclinal and anticlinal divisions during bulge development (beginning a few hours after pollination) than they did during development of the seed coat away from the bulge. Further, the boundary between the square sclereids in the transition zone and the elongated ones of the bulge delineate the edge of the water gap.     

Variation of cleavage pattern permitting normal development in a sand dollar,Peronella japonica: comparison with other sand dollars

 Peronella japonica, a sand dollar, forms an abbreviated pluteus larva and metamorphoses within 3 days without feeding. In the present study, the cleavage pattern of Peronella embryos was found to be quite irregular in the vegetal blastomeres at the fourth cleavage. Less than half of the embryos examined formed four typical micromeres. The majority formed zero, one, two or three typical micromeres of regular size, and the blastomere(s) remaining in the vegetal-most region was atypical in size and/or its direction of division. Most embryos were able to form pluteus larvae and a considerable proportion of these metamorphosed into juvenile sea urchins, regardless of whether or not they had formed four typical micromeres of regular size, although embryos which formed no typical micromeres developed into pluteus larvae less frequently. The micromere progeny in Peronella embryos form skeletogenic mesenchyme cells. The average numbers of skeletogenic mesenchyme cells in the three sand dollar species, Clypeaster japonicus, Astriclypeus manni and P. japonica were 62, 122 and 219, respectively. In these species, the skeletogenic mesenchyme cell-specific glycoprotein (msp130) was first detected immediately after ingression of the primary mesenchyme cells, spicules appeared at the early gastrula stage and triradiate spicules were found in late gastrulae. Appearance of these characteristics was markedly accelerated in the embryos of A. manni and P. japonica in comparison with those of C. japonicus. Each step in the formation of larval spicules was equally accelerated in A. manni and P. japonica, although the appearance of the adult skeleton was further accelerated in P. japonica in comparison with A. manni, possibly because of omission of the four- to eight-armed pluteus stages. Received: 1 September 1995 / Accepted in revised form: 21 May 1995     

Postovipositional development of the sand snake Psammophis sibilans (Serpentes:Lamprophiidae) in comparison with other snake species

Characterizing and comparing developmental progress across different species helps to interpret how different or similar body forms evolved. We present an embryonic table for the oviparous African Sand Snake Psammophis sibilans from the Lamprophidae family, describing its postovipositional in ovo development. Psammophis is a good model of a genus that is widely distributed in Africa and Asia and includes 22 species. We describe ten embryonic stages based on the development of externally visible morphological characteristics such as; pharyngeal arches, facial processes, eyes, scales, body pigmentation and body colour pattern development. This study discusses the development of this snake and compares it with that of the closely related brown house snake Lamprophis fulliginosus (Lamprophidae) and the medically important venomous cobras Naja haje haje and Naja kaouthia from the sister lineage Elapidae. The distantly related basal snake Python sebae, which displays different morphology and behaviour, was chosen for deeper insight into the evolution of body structures within the snake clade. We found interspecific differences in the relative stage of development of embryonic structures at the time of oviposition and during postovipositional embryonic development. One of the outcomes of this study is that embryonic structures such as the pharyngeal processes, eye pigmentation and scales are interspecifically conservative in regard to timing of morphodifferentiation, while body pigmentation, colour and colour pattern are interspecifically plastic in their temporospatial development.     

Changing crop biodiversity and resource use efficiency of traditional versus introduced crops in the cold desert of the northwestern Indian Himalaya: a case of the Lahaul valley

This paper studies the crop biodiversity and its changing patterns across an altitudinal gradient of three highland village agroecosystems: Hinsa (2700 m), Jahlma (3000 m) and Khoksar (3200 m). It also deals with energy (kilocalorie (physiological); kcal or Megajoules; MJ) and monetary (Indian rupee; Rs) efficiency of traditional crops versus recently introduced cash crops in the cold desert of the Lahaul valley. Newly introduced cash crops like hop, pea and potato have drastically reduced the crop biodiversity along the increasing altitudes. Of the total cropped area, 70% is occupied by these cash crops. Organic manure, mainly derived from forests and night soil, accounted for 90 and 77% of the total energy input in traditional and introduced cash crops in all the three selected villages, respectively. The energy output/input for traditional crops ranged in Hinsa from 0.6 (Kuth (Sausurea lappa, family Asteraceae) — a root crop of medicinal use) to 6.1 (amaranths); for the introduced cash crops it varied from 0.9 (potato) in Hinsa to 2.8 (pea) in Jahlma. In terms of energy, traditional crops are richer and more viable than the introduced cash crops. Monetary output/input of traditional crops was 0.8 (maize) to 11.8 (Kuth) and for introduced cash crops it was from 1.8 (potato) to 6.2 (hop). Overall, the average monetary efficiency of introduced crops was higher than that of traditional ones. Taking the average of the three villages, the energy and monetary output/input for the introduced cash crops was 1.4 and 3.3, respectively. However, for the purpose of food security and environmental conservation there is a need to establish a balance between traditional and introduced cash crops. This will make farming sustainable in the cold desert environment where crop growth period is limited. The Lahaul valley is landlocked and due to heavy snow deposits at Rohtang pass (3978 m), its main entry from Kullu valley remains accessible only for 5 months (July–November).     

Interactions between TIME and PIN could play a role in the system that controls the duration of diapause development and synchronization with seasonal cycles in the silkworm Bombyx mori*

The significance of winter cold in the termination of diapause was investigated with regards to TIME and PIN in eggs of the silkworm Bombyx mori. TIME (time interval measuring enzyme) is an ATPase that can measure time intervals by exhibiting a transitory burst of activation of the enzyme in accordance with diapause development, which requires cold for resumption of embryonic development in the silkworm. The possible timer function of TIME comprises a built‐in mechanism in the protein structure. TIME is a metallo‐glycoprotein consisting of 156 amino acid residues with a unique sequence in the N‐terminal region to which a sugar chain is attached. PIN (peptidyl inhibitory needle) inhibits the ATPase activity of TIME. PIN is not a simple enzyme inhibitor, but holds the timer by forming a time‐regulatory complex with TIME. The carbohydrate moiety of TIME is essential for the assembly of a high‐affinity PIN‐binding site within the timer motif of the TIME structure. The binding interaction between TIME and PIN was much tighter (nearly 1000 times) at 25°C than that at 4°C, as measured by fluorescence polarization. Because the logEC₅₀ at 4°C was approximately 7 nmol/L, PIN must dissociate from TIME at the physiological concentration of TIME in eggs in the winter cold. Based on the results of our study, we propose that the dissociation of the TIME–PIN complex in the winter cold cues a series of conformational changes of TIME, ultimately reaching the active form of ATPase which in turn causes the completion of diapause development and initiates new developmental programs.