种子顽拗性:最新评价

根据种子的脱水行为,可以把种子分为正常性、顽拗性和中间性种子三种类型.种子脱水耐性和脱水敏感性的鉴定是制定种子的贮藏策略和物种基因资源长期保存的依据.除了物种的内在特性外,种子的发育状态、脱水速率、脱水时和重新水合时的环境是影响种子脱水耐性的重要因子.种子的存活率、电解质渗漏速率和存活种子的萌发生长速率是衡量种子脱水耐性的良好的综合参数.种子的脱水耐性是一种数量性状,用"临界含水量"的概念来判断是不正确的,且在评价种子顽拗性中引起了一些混乱.本文还提出了一种全新的评价种子顽拗性的工作模式.     

种子顽拗性：最新评价

根据种子的脱水行为,可以把种子分为正常性、顽拗性和中间性种子三种类型。种子脱水耐性和脱水敏感性的鉴定是制定种子的贮藏策略和物种基因资源长期保存的依据。除了物种的内在特性外,种子的发育状态、脱水速率、脱水时和重新水合时的环境是影响种子脱水耐性的重要因子。种子的存活率、电解质渗漏速率和存活种子的萌发生长速率是衡量种子脱水耐性的良好的综合参数。种子的脱水耐性是一种数量性状,用“临界含水量”的概念来判断是不正确的,且在评价种子顽拗性中引起了一些混乱。本文还提出了一种全新的评价种子顽拗性的工作模式。     

顽拗性种子研究

Roberts（1973）根据种子的贮藏行为把种子分为正常性种子和顽拗性种子,Ellis（1990）等认为红这两者之间还存在中间类型即中间型种子。种子的顽拗性是一个数量性状,具有连续性：分为低度顽拗性种子、中度顽拗性种子、高度顽拗性种子。具顽拗性种子的植物主要来自两类：一类为水生植物,如：茭白、菱等;另一类主要是具大粒种子的多年生木本,包括热带作物,热带水果,热带林木,如：可可,红毛丹等。     

种子顽拗性的形成机理及其保存技术

根据种子的脱水行为将种子分为正常性种子、顽拗性种子和中间性种子。顽拗性种子在发育的末期不经历成熟脱水．脱落时有相对高的含水量,并且对低温和脱水干燥非常敏感。在自然界,顽拗性种子存在一个连续群,即低度、中度和高度顽拗性种子,其差异在于对脱水伤害的敏感程度。影响种子顽拗性的因素,既有种子本身的生理生化物质基础,也有种子在母株上发育过程中所经受的外界环境的影响。目前,对种子脱水耐性的分子机制及其保存技术研究得较多。本文综述了有关顽拗性种子研究的近期进展。     

顽拗型种子的生物学特性及种子顽拗性的进化

综述了顽拗型种子的形态、大小、含水量、植物分类、植物生态方面的一般特性,分析了顽拗型种子对环境的生态适应性,并讨论了种子顽拗性的可能进化模式,进而指出顽拗型种子生理生态学研究的意义和应用前景.顽拗型种子一般千粒重和体积较大,含水量较高,萌发迅速且多无休眠特性;产生顽拗性种子的植物分布很广,与其系统分类地位无关,但多起源于湿润的生境;目前尚无足够的证据表明种子顽拗性是原始性状或是衍生性状,要解决这一问题还需更深入的研究,尤其是种子生理学和生态学家的合作研究.     

顽拗性种子

本文讨论了顽拗性种子的基本特性,在劣变中的生理生化变化,脱水敏感性的细胞生物学基础,以及贮藏顽拗性种子的必要性和可能性。     

植物顽拗性种子研究进展

对植物顽拗性种子的概念、物种分类、形态、分布及一些生理生态特征进行了综述,分析了顽拗性种子脱水敏感性的原因和对环境的生态适应性,探讨其可能的进化地位和贮存技术。同时,对顽拗性种子研究的发展趋势和应用前景进行了展望,即加强生境调查,更新顽拗性种子植物数据库,从形态学、生理生态学和分子生物学分析种子顽拗性本质。     

植物种质资源超低温保存概述

简要回顾了植物种质资源超低温保存的历史,说明了超低温保存植物材料的多样性,阐述了超低温耐性的生物学基础及超低温伤害产生的原因和类型,介绍了各种常用超低温保存方法的技术要点,并对生产顽拗性种子的植物种质资源的超低温保存作了专门的论述,分析了生产顽拗性种子的植物种质资源超低温保存的潜力、现状和困难,指出顽拗性种子的超低温保存是植物种质资源超低温保存的重点和难点,而真正实现用超低温保存技术贮藏顽拗性植物种质资源还有很长的路要走。     

低温保存技术在顽拗性种子种质保存中的利用

由于顽拗性种子不耐脱水且对低温敏感,常规保存方法难以达到长期保存的目的。因此,(超)低温保存顽拗性种子种质是最理想的方法。顽拗性种子的低温保存,应用较多的是玻璃化法和两步法。诸多因素影响着低温保存的成败,如种子或胚的含水量水平、溶液低温保护剂效应、降温冰冻与解冻方式、水合过程以及后培养等,这些需深入探索与解决。除顽拗性种子脱水耐性和低温敏感性机理外,植物细胞的冻害和抗冻机理也亟需探明,以便找到最佳冷冻方法,制定长期保存种质基因的最佳方案。     

顽拗性种子的研究进展

根据种子的贮藏特性,Roberts将种子分为正常性种子(Orthodox seeds)和顽拗性种子(Recalcitrant seeds)两大类。正常性种子包括大多数农作物种子,能够忍受脱水和低温,在5±1％含水量和-18℃低温下贮藏,种子的生活力可保持相当一段时间,有的可达数十年甚至更长,在我国东北地区天然干涸的湖床中发现的印度莲子是很老的活种子,应用放射性碳测定,据称这些种子的寿命在1000年以上。     

From Avicennia to Zizania: seed recalcitrance in perspective

BACKGROUND: Considered only in terms of tolerance of, or sensitivity to, desiccation (which is an oversimplification), orthodox seeds are those which tolerate dehydration and are storable in this condition, while highly recalcitrant seeds are damaged by loss of only a small proportion of water and are unstorable for practical purposes. Between these extremes, however, there may be a gradation of the responses to dehydration--and also to other factors--suggesting perhaps that seed behaviour might be best considered as constituting a continuum subtended by extreme orthodoxy and the highest degree of recalcitrance. As the characteristics of seeds of an increasing number of species are elucidated, non-orthodox seed behaviour is emerging as considerably more commonplace--and its basis far more complex--than previously suspected. SCOPE: Whatever the post-harvest responses of seeds of individual species may be, they are the outcome of the properties of pre-shedding development, and a full understanding of the subtleties of various degrees of non-orthodox behaviour must await the identification of, and interaction among, all the factors conferring extreme orthodoxy. Appreciation of the phenomenon of recalcitrance is confounded by intra- and interseasonal variability across species, as well as within individual species. However, recent evidence suggests that provenance is a pivotal factor in determining the degree of recalcitrant behaviour exhibited by seeds of individual species. Non-orthodox--and, in particular, recalcitrant--seed behaviour is not merely a matter of desiccation sensitivity: the primary basis is that the seeds are actively metabolic when they are shed, in contrast to orthodox types which are quiescent. This affects all aspects of the handling and storage of recalcitrant seeds. In the short to medium term, recalcitrant seeds should be stored in as hydrated a condition as when they are shed, and at the lowest temperature not diminishing vigour or viability. Such hydrated storage has attendant problems of fungal proliferation which, unless minimized, will inevitably and significantly affect seed quality. The life span of seeds in hydrated storage even under the best conditions is variable among species, but is curtailed (days to months), and various approaches attempting to extend non-orthodox seed longevity are discussed. Conservation of the genetic resources by means other than seed storage is then briefly considered, with detail on the potential for, and difficulties with, cryostorage highlighted. CONCLUSIONS: There appears to be little taxonomic relationship among species exhibiting the phenomenon of seed recalcitrance, suggesting that it is a derived trait, with tolerance having been lost a number of times. Although recalcitrant seededness is best represented in the mesic tropics, particularly among rainforest climax species, it does occur in cooler, drier and markedly seasonal habitats. The selective advantages of the trait are considered.     

Trends in the evolution of dicotyledonous seeds based on character associations, with special reference to pachychalazy and recalcitrance

VON TEICHMAN, I. & VAN WYK, A. E., 1991. Trends in the evolution of dicotyledonous seeds based on character associations, with special reference to pachychalazy and recalcitrance. The possible evolutionary status of the endothelium, hypostase, pachychalaza and the recalcitrant viability behaviour of seeds is considered in relation to bitegmy/unitegmy, crassinucellate/tenuinucellate ovules, nuclear/cellular endosperm development, large/small seed size, woody/herbaceous habit and tropical/temperate habitat. The presence of the endothelium, hypostase, pachychalaza and recalcitrance in dicotyledonous families is plotted against Dahlgren's system of classification. Results are compared with Sporne's advancement index for the various families. An endothelium is considered derived since it occurs more often in highly evolved superorders and is significantly associated with derived ovule and endosperm character states as well as with smaller seed size. A hypostase appears to be relatively ancestral and is significantly associated with pachychalazy and recalcitrance. The endothelium and hypostase have developed independently in many taxa and could be interpreted as being structurally and functionally analogous. Pachychalazy and recalcitrance are significantly associated with ancestral ovule character states and, at the species level, with large seed size (overgrown seed), woody habit and tropical habitat. The presence of pachychalazy, recalcitrance and associated large seed size are therefore regarded as ancestral character states of the dicotyledons. Consideration of currently accepted dicta on seed character state polarity, suggests a reversal in the evolutionary status of pachychalazy and large seed size.     

Comparative analysis of metabolome of rice seeds at three developmental stages using a recombinant inbred line population

Plants are considered an important food and nutrition source for humans. Despite advances in plant seed metabolomics, knowledge about the genetic and molecular bases of rice seed metabolomes at different developmental stages is still limited. Here, using Zhenshan 97 (ZS97) and Minghui 63 (MH63), we performed a widely targeted metabolic profiling in seeds during grain filling, mature seeds and germinating seeds. The diversity between MH63 and ZS97 was characterized in terms of the content of metabolites and the metabolic shifting across developmental stages. Taking advantage of the ultra‐high‐density genetic map of a population of 210 recombinant inbred lines (RILs) derived from a cross between ZS97 and MH63, we identified 4681 putative metabolic quantitative trait loci (mQTLs) in seeds across the three stages. Further analysis of the mQTLs for the codetected metabolites across the three stages revealed that the genetic regulation of metabolite accumulation was closely related to developmental stage. Using in silico analyses, we characterized 35 candidate genes responsible for 30 structurally identified or annotated compounds, among which LOC_Os07g04970 and LOC_Os06g03990 were identified to be responsible for feruloylserotonin and l ‐asparagine content variation across populations, respectively. Metabolite?agronomic trait association and colocation between mQTLs and phenotypic quantitative trait loci (pQTLs) revealed the complexity of the metabolite?agronomic trait relationship and the corresponding genetic basis.     

Recalcitrant Seeds: Short-term Storage Effects in Avicennia marina (Forsk.) Vierh. may be Germination-associated

When recalcitrant propagules (seeds) of Avicennia marina werestored in a dry air stream there was no significant change intheir moisture content for 10 days, after which it declinedrapidly. Seed viability remained high during this 10-d period,only then declining. Ultrastructural changes in embryonic rootprimordium cells during the 10-d storage period were similarto those characterizing germination. It is suggested that thecontinuation of germination processes after shedding may beresponsible for the deterioration in stored recalcitrant seeds. Avicennia marina, recalcitrance, seed storage, germination     

Dynamics of Water and Abscisic Acid During Embryogeny and Embryo Drying in the Recalcitrant Seeds of Vateria indica L.

Vateria indica L. is a critically endangered tree species in South-Western Ghats of India, commercially exploited for its valuable resins. Seed recalcitrance is a major problem hindering the natural regeneration of this species and it poses a great challenge in seed storage and conservation. There was a continuous import of water from the maternal tissues to seed tissues till maturity and the seeds were released in a fully hydrated state. Differential accumulation of water has been noticed in the cotyledons and embryonal axis. There was a positive correlation between seed moisture content and rate of germination which is a character of recalcitrant seeds. The critical moisture content was found to be 40% in the axis and 23.5% in the cotyledons, below which the embryo will not germinate. Loss of germination ability as a result of desiccation was attributed to the cell membrane damage, expressed as the electrolyte leakage exceeding 0.79 μS/cm. ABA peaked in the mid embryogenesis, then dropped drastically and maintained a lower level till seed maturity. On desiccation, ABA started to increase but gradually dropped down. Both cotyledons and embryonal axis had differential ABA content but exhibited a general pattern of ABA level during embryogeny. Due to the thin seed coat/embryo ratio and low investment in the seed coat, this recalcitrant seed could not hold water as efficient as orthodox seeds. Thus, it germinated as soon as it was shed from the mother plant. On desiccation, ABA shot up and moisture content decreased along with electrolyte leakage and cell membrane damage. All these hindered germination of the seed. Thus, we can see a clear interplay between moisture content and ABA levels during embryogeny and desiccation. Since the seed biology of this species has not been well documented, the present work is mainly intended to study the dynamics of water and ABA during embryogeny and embryo drying. This study can surely contribute to the long-term storage and conservation of recalcitrant seeds which is a less explored area.

     

Inheritance of seed desiccation sensitivity in a coffee interspecific cross: evidence for polygenic determinism

The genetic determinism of seed desiccation sensitivity was studied using a cross between two coffee species exhibiting a large difference for this trait, Coffea pseudozanguebariae (tolerant) and C. liberica (sensitive). Throughout the whole study, seed desiccation tolerance was quantified both in terms of water content and water activity. Whatever the parameter used, the level of seed desiccation tolerance in F1 hybrids corresponded to that of the mid-parent, thus indicating an additive inheritance of seed desiccation tolerance at the F1 level. A broad variation was observed among hybrids backcrossed to C. liberica (BCs) for seed desiccation tolerance, independent of the parameter used to quantify it. This variation was continuous and BCs showed transgression in the direction of the most desiccation sensitive parent, indicating (i) that desiccation tolerance is a polygenic trait in coffee species, and (ii) that C. pseudozanguebariae does not present the most favourable alleles for all the genes involved in seed desiccation tolerance. No significant difference was observed between the two reciprocal backcrosses, F1xC. liberica and C. libericaxF1, for the level of desiccation tolerance of their seeds, showing the absence of a maternal effect on this trait. There was no significant effect of the number of seeds harvested from each BC on the level of desiccation tolerance of its seeds. Moreover, there was no significant correlation within BCs between seed size, seed viability, and water content before desiccation and desiccation tolerance.     

Responses to Drying of Recalcitrant Seeds of Quercus nigra L.

It has been suggested that rate of desiccation can influencethe expression of recalcitrant behaviour in seeds, thus complicatingthe task of determining which seeds are truly recalcitrant.The objective of this study was to see if variable rates ofdesiccation influenced such behaviour inQuercus nigra L., atree seed known to be recalcitrant. Acorn moisture content, leachate conductivity, and germinationwere determined at various times during desiccation at threerates at 27 and 40°C. Moisture contents and germinationdecreased as the severity of desiccation increased. Leachateconductivity increased slightly but was not a sensitive indicatorof loss of viability. The critical (lethal) moisture contentfor these acorns was 10–15%, although rehydration within48 h of reaching this level prevented death in about 25% ofthe acorns. At 27°C any desiccation treatment that producedlosses of 30–50mg of moisture per g of acorn dry weightper day should be suitable as a test for recalcitrance in thegenus. Apparent physiological or fungal damage at 40°C rulesout the higher temperature for such a test. Quercus nigra L.; desiccation rate; temperature; recalcitrant; leachate conductivity; germination; viability     

Dormancy Breaking and Storage Behavior of Garcinia cowa Roxb. (Guttiferae) Seeds: Implications for Ecological Function and Germplasm Conservation

The dormancy breaking and storage behavior of Garcinia cowa Roxb. seeds were investigated.The seeds of G. cowa had 8-11 months dormancy in their natural habitat. Seeds were matured and dispersed at the end of the rainy season (mid-late August to late September) and were scatter-hoarded by rodents as food for winter after the seeds had fallen to the ground. Seedlings often emerged in the forest during the rainy season (May to August) the following year. Intact seeds of G. cowa failed to germinate after being sown at 30 ℃ for 120 d and the mean germination time (MGT) of seeds cultured in a shade (50% sunlight)nursery was 252 d. The most effective method of breaking dormancy was to remove the seed coat totally,which reduced the MGT to 13 d at 30 ℃. Germination was also promoted by partial removal of the seed coat (excising the hilum and exposing the radicle) and chemical scarification (immersion in 1% H2O2 for 1 d).Unscarified seeds take up water rapidly in the first 96 h, but water was absorbed by the outside seed coat,without penetrating through it. The moisture content (MC) of G. cowa seeds was high (50% in fresh weight)at shedding. The seeds could tolerate desiccation to some extent, until the MC reached approximately 40%;below that, the viability decreases rapidly and all seeds died at approximately 17% of MC. Seed viability decreased rapidly when seeds were chilled at 4 ℃; germination was 2% after storage for 1 week. Even stored at 10 ℃, seeds began to be damaged after 4 weeks. Seed storage for 1 yr revealed that in both dry (relative humidity (35 ± 5)%) and moist (wet sand) storage conditions, seed viability declined, but germination percentages for seeds stored under moist conditions are better than for seed stored under dry conditions.Because of their low tolerance to desiccation, marked chilling sensitivity and relatively short lifespan, G.cowa seeds should be classified into the tropical recalcitrant category. The ecological implications of dormant recalcitrant seeds and cues on storing recalcitrant seeds were discussed.     

拟南芥低温黑暗发芽基因的鉴定及QTL定位

低温发芽是一个重要农艺性状,提高作物的耐寒性并确定控制低温发芽的基因是一个全球性的重大育种课题。通过对300多份拟南芥种子进行发芽试验,鉴定了低温黑暗下发芽的自然变异,并比较了两种生态型Bay-0和Shahdara低温黑暗下的发芽特性,这种自然变异被用来鉴定低温黑暗中发芽基因的遗传位点;利用这两个生态型杂交产生的重组近交系,结合分子标记和QTL方法,检测到种子在低温黑暗下发芽至少受6个QTL所控制,其中3个为主效位点,它们所解释的自然变异为61%,利用异源近亲代(HIF)验证了这3个主要QTL,为低温黑暗发芽基因的精细定位和基因克隆等研究奠定了基础。