栽培稻抗旱性研究的现状与策略

 水资源短缺正成为制约我国农业发展的重要因素。培育抗旱的栽培稻品种并实现水稻旱作，不但可在很大程度上节约水资源，而且有利于增产稳产，节约能源和减少环境污染。抗旱性包括逃旱性、避旱性、耐旱性和复原抗旱性。形态生理学的研究揭示出大量的与栽培稻抗旱性有关的形态特征和生理特性，如根系和叶片性状、生育期、渗透调节、脱落酸含量与栽培稻抗旱性密切相关，且已利用分子标记对上述性状进行了基因定位（QTL）研究。旱稻品种改良也已取得重大进展。在进行抗旱品种改良的基础上，通过引进相应的栽培技术，节水种植，实现水稻旱作，并达到稳产与增产的目的，是抗旱性研究的战略目标。在增产、稳产和优质的前提下，以培育耐旱性极强的水稻（或旱稻）为中心，建立有代表性的抗旱性研究基地, 进一步加强稻属抗旱基因资源的发掘和创新、抗旱生理学和遗传学的研究、利用现代生物技术实现不同物种间抗旱基因的转移、建立节水种植栽培技术新体系是目前抗旱性研究的主要内容。     

水稻生理特性与抗旱性的相关分析及QTL定位

利用籼稻品种IR64和粳稻品种Azucena杂交产生的包含110个加倍单倍体株系的群体,在干旱胁迫和正常水分条件下,连续在2004年和2005年于抽穗期分别测定了叶片水势、相对含水量、叶绿素含量(SPAD值)、游离脯氨酸含量、气孔导度和蒸腾速率,并于成熟期取样,计算抗旱系数。与正常水分状况下相比,干旱胁迫条件下叶片的游离脯氨酸含量的增加达极显著水平,干旱胁迫条件下叶片的相对含水量、水势、叶绿素含量和气孔导度的降低均达显著或极显著水平。相关分析表明,在干旱胁迫条件下,叶片相对含水量、叶片水势与抗旱系数呈显著或极显著正相关。 利用175个RFLP标记构建的遗传连锁图谱分析了与抗旱性相关的叶片生理指标,共检测到与抗旱性相关的6个生理指标的7个加性QTL,31对上位性QTL,其中有2个主效QTL、9对上位性QTL存在环境互作效应。在两种水分条件下检测到的QTL结果有较大差异,说明干旱胁迫对控制与抗旱性相关的叶片生理性状基因的表达有显著的影响。在6个抗旱相关生理指标中,检测到的控制叶片气孔导度和水势的QTL较多,有3个加性QTL和8对上位性QTL控制气孔导度,有8对上位性QTL控制水势。     

Mapping quantitative trait loci associated with root growth in upland rice (Oryza sativa L.) exposed to soil water-deficit in fields with contrasting soil properties

A mapping population of 114 lines from Bala × Azucena was grown under drought stress at two field sites with contrasting soil physical properties. Drought was imposed between 35 and 65 days after sowing (DAS) and root density at 35 cm depth was measured 70 DAS. Leaf rolling, leaf drying and relative water content were recorded as indicators of drought avoidance. Root density correlated with indicators of drought avoidance. Two significant and two putative quantitative trait loci (QTLs) for root density and 28 QTLs for drought avoidance were identified. Most QTLs did not agree between sites. There was also reasonable agreement between leaf-drying QTLs and previously reported root-growth QTLs detected under controlled conditions (in contrast to a previous screen on soil with a higher penetration resistance). These data also reveal QTL × environment interaction, which will need to be understood more clearly if progress towards breeding for drought resistance via alterations of root morphology is to be achieved.     

Genetic analysis and validation of quantitative trait loci associated with reproductive-growth traits and grain yield under drought stress in a doubled haploid line population of rice (Oryza sativa L.)

Drought is a major constraint for rice production and yield stability in rainfed ecosystems, especially when it occurs during the reproductive stage. Combined genetic and physiological analysis of reproductive-growth traits and their effects on yield and yield components under drought stress is important for dissecting the biological bases of drought resistance and for rice yield improvement in water-limited environments. A subset of a doubled haploid (DH) line population of CT9993-5-10-1-M/IR62266-42-6-2 was evaluated for variation in plant water status, phenology, reproductive-growth traits, yield and yield components under reproductive-stage drought stress and irrigated (non-stress) conditions in the field. Since this DH line population was previously used in extensive quantitative trait loci (QTLs) mapping of various drought resistance component traits, we aimed at identifying QTLs for specific reproductive-growth and yield traits and also to validate the consensus QTLs identified earlier in these DH lines using meta-analysis. DH lines showed significant variation for plant water status, reproductive-growth traits, yield and yield components under drought stress. Total dry matter, number of panicles per plant, harvest index, panicle harvest index, panicle fertility, pollen fertility, spikelet fertility and hundred grain weight had significant positive correlations with grain yield under drought stress. A total of 46 QTLs were identified for the various traits under stress and non-stress conditions with phenotypic effect ranging from 9.5 to 35.6% in this study. QTLs for panicle exsertion, peduncle length and pollen fertility, identified for the first time in this study, could be useful in marker-assisted breeding (MAB) for drought resistance in rice. A total of 97 QTLs linked to plant growth, phenology, reproductive-growth traits, yield and its components under non-stress and drought stress, identified in this study as well as from earlier published information, were subjected to meta-analysis. Meta-analysis identified 23 MQTLs linked to plant phenology and production traits under stress conditions. Among them, four MQTLs viz., 1.3 for plant height, 3.1 for days to flowering, 8.1 for days to flowering or delay in flowering and 9.1 for days to flowering are true QTLs. Consensus QTLs for reproductive-growth traits and grain yield under drought stress have been identified on chromosomes 1 and 9 using meta-QTL analysis in these DH lines. These MQTLs associated with reproductive-growth, grain yield and its component traits under drought stress could be useful targets for drought resistance improvement in rice through MAB and/or map-based positional analysis of candidate genes.     

Upland rice grown in soil-filled chambers and exposed to contrasting water-deficit regimes: II. Mapping quantitative trait loci for root morphology and distribution

Root morphological characteristics are known to be important in the drought resistance of some rice (Oryza sativa L.) varieties. The identification of quantitative trait loci (QTLs) associated with root morphology and other drought resistance-related traits should help breeders produce more drought resistant varieties. Stability in the expression of root growth QTL across rooting environments is critical for their use in breeding programs. A greenhouse experiment in which a mapping population of 140 recombinant inbred lines and the parental varieties Bala and Azucena were grown in glass-sided soil chambers and evaluated for root growth and water uptake was conducted. In each of 2 years, two treatments were used; an early water-deficit (WD0) in which seeds were sown into wet soil but received no more water, and a late water-deficit (WD49) in which the plants were watered for 49 days and then received no water for a week. The major differences between treatments and years in dry matter partitioning and root growth traits are reported elsewhere. Here, the identification of QTLs for root growth traits by composite interval mapping is described. At LOD>3.2, there were six QTLs for the weight of roots below 90 cm and maximum root length, 11 for root to shoot ratio, 12 for the number of roots past 100 cm, and 14 for root thickness. A total of 24 regions were identified as containing QTLs (these regions often contained several QTLs identified for different root traits). Some were revealed only in individual experiments and/or for individual traits, while others were common to different traits or experiments. Seven QTLs, on chromosomes 1, 2, 4, 7, 9 (two QTLs) and 11, where considered particularly noteworthy. The complex results are discussed in the context of previously reported QTLs for root growth in other populations, the interaction between QTL with the environment and the value of QTLs for breeding.     

Inheritance and QTL Mapping of Salt Tolerance in Rice

An F2 population derived from the cross between Jiucaiqing (japonica) and IR36 (indica) was used to analyze the inheritance of salt tolerance in rice by genetic model of major-genes plus polygenes, and to map the corresponding QTLs by SSR molecular markers. Rice plants of P1, P2, F1 and F2 at 5- to 6- leaf stage were treated under 140 mmol/L NaCI for 10 days. Three indices representing the ability of salt tolerance of rice seedlings were measured, including salt tolerance rating (STR), Na^ /K^ ratio in roots and dry matter weight of shoots (DWS). STR, Na^ /K^ and DWS were all controlled by two major genes with modification by polygenes. Heritability of these traits from major genes was 17.8, 53.3 and 52.3%, respectively. The linkage map constructed by 62 SSR molecular markers covered a total length of about 1 142 cM. There were three QTLs detected for STR located on chromosome 1, 5 and 9, two QTLs for DWS on chromosomes 8 and 9, and two QTLs for Na^ /K^ on chromosomes 2 and 6, one on each chromosome respectively. Single QTL accounted for 6.7 to 19.3% of phenotypic variation. Identification method of salt tolerance in rice and breeding of rice varieties with salt tolerance based on molecular markers assisted selection had been discussed.     

Rice Root Genetic Architecture: Meta-analysis from a Drought QTL Database

During the last 10 years, a large number of quantitative trait loci (QTLs) controlling rice root morphological parameters have been detected in several mapping populations by teams interested in improving drought resistance in rice. Compiling these data could be extremely helpful in identifying candidate genes by positioning consensus QTLs with more precision through meta-QTL analysis. We extracted information from 24 published papers on QTLs controlling 29 root parameters including root number, maximum root length, root thickness, root/shoot ratio, and root penetration index. A web-accessible database of 675 root QTLs detected in 12 populations was constructed. This database includes also all QTLs for drought resistance traits in rice published between 1995 and 2007. The physical position on the pseudo-chromosomes of the markers flanking each QTL was determined. An overview of the number of root QTLs in 5-Mb segments covering the whole genome revealed the existence of “hot spots,” The 32 trait × chromosome combinations comprising six or more QTLs were subjected to a meta-QTL analysis using the software package MetaQTL. The method enabled us both to determine the likely number of true QTLs in these areas using an Akaike information criterion and to estimate their position. The meta-QTL confidence intervals were notably reduced and, for the smallest ones, encompassed only a few genes.     

六棱大麦抗赤霉病QTL的定位(英文)

赤霉病是最严重的大麦病害之一。由于赤霉病抗性是受多基因控制的数量性状(QTL),并且一些表型性状也影响大麦赤霉病的抗病,如棱数、株高和抽穗期等,所以抗赤霉病大麦品种的选育十分困难。为了明确加拿大六棱大麦中赤霉病抗性以及相关性状的QTLs,本研究在4年中对93个家系的DH作图群体中赤霉病抗性、呕吐毒素(DON)含量、株高、抽穗期和成熟期等相关性状进行调查,并利用分子标记(444个DArT和26个SSR标记)构建的连锁图谱对QTL开展复合区间作图。结果表明,本研究共检测到4个影响赤霉病的QTLs,其中,2个主要的QTLs定位在3H和7H染色体上,它们的加性效应为-3.44和-3.69,分别解释14.1%和17.5%的表型差异,总共解释31.6%的赤霉病抗性差异;另外2个QTLs定位于7H染色体上,但二者同时也与DON含量显著相关。此外,在3H、5H和7H染色体上确定了5个影响株高的QTLs,在2H、4H、5H和7H上确定了4个影响抽穗期的QTLs。同时发现2个赤霉病抗性QTLs和1个DON累积QTL与控制株高的QTLs聚集重叠,1个赤霉病抗性QTL和抽穗期QTLs重叠。这些与赤霉病抗性、株高及抽穗期等农艺性状紧密连锁的分子标记可进一步用于有效提高抗赤霉病大麦品种的选育效率。     

土壤干旱胁迫对不同水稻品种叶片卷曲的影响

在人工控制土壤水分条件下,以13个具有不同耐旱性的栽培稻品种（组合）为供试材料,研究了在分蘖盛期土壤断水后随着干旱胁迫的加重叶片卷曲度（LRI）的动态变化及其与蒸腾、光合特性和叶片水分利用的关系。结果表明,不同品种的LRI明显增大的起始日期、明显增大期日均增量和最大值存在着显著差异,并依据叶片卷曲表现可将参试品种划分为敏感型（如中旱1号）、中间型（如旱稻8号）和钝感型（如巴西陆稻）三种类型。随着土壤断水后干旱胁迫的加重,LRI逐渐增大。LRI与气孔导度（SC）、蒸腾速率（TR）和净光合速率（Pn）的关系因品种类型而异;LRI每增加1级,中旱1号、旱稻8号和巴西陆稻的SC分别降低0.26、0.20和0.17 mm/s,TR分别降低0.86、0.66和0.49 mmol/(m2·s);Pn呈显著下降的叶片卷曲度中旱1号明显小于巴西陆稻。当叶片适度卷曲时,WUE随LRI值的增大而提高,中旱1号LRI的变化同时引起TR和Pn显著改变进而影响WUE,而旱稻8号和巴西陆稻LRI变化对WUE的影响主要系TR改变所致。     

小麦叶片蜡质含量与水分利用效率和产量的关系

为了进一步了解植物叶片蜡质含量与植物抗旱节水性的关系，本试验对6个小麦品种在灌浆后期的旗叶蜡质含量与抗早生理指标进行了相关分析，结果表明，干早条件下小麦灌浆后期叶片蜡质含量与光合速率、叶温、叶片水分利用效率和产量呈显著正相关．蜡质含量高的品种叶片温度高、蒸腾速率大，这与传统现点不同．分析认为蜡质含量高的品种在干早胁迫条件下可能叶片细胞膜稳定性强，气孔不容易发生关闭，因此其叶片水分利用效率和产量较高．蜡质含量除了受遗传因素决定外，还受环境因素影响．     

利用染色体片段置换系群体检测水稻叶片形态QTL

 水稻叶片的形态改良是水稻株型育种和产量育种的重要目标之一。以9311/日本晴染色体片段置换系（CSSLs）群体为材料,定位了上3叶叶片长、宽、叶面积共9个性状QTL,分析了叶片性状与产量性状之间的相关性,同时定位了主穗重及产量构成因素（颖花数、千粒重、结实率）相关QTL。结果表明,CSSLs群体的叶片性状之间存在显著或极显著相关性;叶片性状与主穗重呈显著或极显著正相关,与主穗颖花数呈极显著正相关;叶片形态多数性状与结实率、千粒重没有显著相关性。两年共定位到20个叶片性状QTL,分布于第1、3、4、5、6、9、11共7条染色体的10个区间,贡献率为3.82%~14.61%,其中贡献率大于10%有6个,多个QTL成簇分布在相同区间,3个QTL在两年间重复检测到,8个QTL为前人未报道的新位点。两年共检测到16个与控制主穗产量相关的QTL,分布于第1、2、3、5、7、8、10共7条染色体13个区间,其中有7个主穗产量相关QTL所在5个区间与叶片形态14个QTL所在区间一致。     

The potential use of plant physiological responses to water stress as an indication of varietal sensitivity to drought in four potato (Solanum tuberosum L.) varieties

Four potato varieties were subjected to water stress under controlled conditions. Leaf relative water content, leaf diffusive resistance, and photosynthesis were measured on stressed and unstressed plants during a stress period and during the recovery period following stress relief. Subsequent to the stress, plants were grown to maturity with optimal water supply. Mature plants were harvested and tuber yield and haulm production measured. Stressed plants of all varieties exhibited an increase in leaf diffusive resistance and a decrease in relative water content, transpiration, and photosynthesis as soil moisture decreased. Significant varietal differences in leaf diffusive resistance of stressed plants are apparent and offer promise for development of a screening technique for varietal sensitivity to drought based on stomatal response to water deficits.     

Rice near-isogenic-lines (NILs) contrasting for grain yield under lowland drought stress

The development of near-isogenic-lines (NILs) is a very important tool for both genetic and physiological dissection of drought resistance in rice. Two pairs of NILs differing for grain yield under drought stress were isolated and characterized for yield, yield related traits, and several physiological traits in a range of contrasting environments. In replicated field trials both NIL pairs differed significantly for grain yield under drought stress but showed similar yield potential, phenology, and yield component traits under non-stress conditions. A polymorphism analysis study with 491 SSRs revealed that both NIL pairs are at least 96% genetically similar. These NILs show that small genetic differences can cause large difference in grain yield under drought stress in rice. In both pairs the drought-tolerant NILs showed a significantly higher assimilation rate at later stages both under stress and non-stress conditions. They also had a higher transpiration rate under non-stress condition. The most tolerant NIL (IR77298-14-1-2-B-10) had significantly higher transpiration rate and stomatal conductance in severe stress conditions. In one pair the tolerant NIL had constitutively deeper roots than the susceptible NIL. In the second pair, which had higher mean root length than the first pair, the tolerant NIL had more roots, greater root thickness, and greater root dry weight than the susceptible NIL. Deeper root length may allow tolerant NILs to extract more water at deeper soil layers. It is concluded that enhanced rooting depth is an important strategy for dehydration avoidance and rice adaptation to drought stress, but root architecture might not be the only mechanism causing the significant yield increase we observed in lowland drought stress environments. To further dissect the drought avoidance mechanisms in rice, analysis of root hydraulic properties may be necessary.     

利用重测序的水稻染色体片段代换系群体定位剑叶形态QTL

 剑叶形态性状（剑叶长、剑叶宽和剑叶面积）是水稻理想株型育种的重要目标性状之一。发掘新的控制水稻剑叶形态性状的基因资源,准确鉴定和定位水稻剑叶形态性状QTL,对开展水稻剑叶形态性状分子生物学研究和理想株型分子育种都具有重要意义。 以通过高通量测序而准确获知代换片段位置及长度的一套用籼稻品种9311为受体、粳稻品种日本晴为供体构建的,包括128个染色体片段代换系群体为材料,对剑叶形态性状及其与主穗颖花之间的相关性进行分析,结果表明剑叶面积与剑叶长、宽呈极显著正相关,主穗颖花数与剑叶长、剑叶面积呈极显著正相关。利用多元回归分析方法,结合Bin map,共鉴定出与水稻剑叶长、宽和面积相关的QTL分别为4、4和6个,贡献率介于4.08%~60.40%。上述QTL的准确定位,为进一步精细定位及克隆相应QTL以及开展水稻剑叶形态性状分子育种奠定了基础。     

Faba bean breeding for drought-affected environments: A physiological and agronomic perspective

The faba bean (Vicia faba L.) crop often experiences drought during its growth and development such that soil moisture deficits constrain its production. As droughts are predicted to increase in both frequency and intensity due to climate change, a better understanding of drought response patterns and associated traits is essential for obtaining yield stability in water-limited environments. This review deals with adaptation mechanisms associated with drought avoidance, escape and tolerance, with an emphasis on physiological traits such as stomatal conductance, carbon isotope discrimination and leaf temperature. Leaf temperature is considered an effective surrogate measure for other measures of stomatal characteristics. Drought tolerance through osmotic adjustment has not yet been demonstrated in faba bean although it is found in many other legumes including chickpea and pea. Deeper root growth, leading to uptake of otherwise unavailable water, helps the plant to avoid drought by delaying dehydration, but genetic variation and heritability of the trait are essentially unknown for faba bean. Crop management strategies, such as early planting, and appropriate phenology, are particularly important for drought escape in regions where terminal drought is common. Disease resistance is especially important in drought-prone areas to reduce the need for expensive control measures when yields are uncertain. The relevance of soil fertility status and nutrient availability are also covered. Drought escape and ascochyta blight resistance are important breeding objectives for terminal drought regions. Some form of drought resistance is necessary for the transient droughts experienced in most regions, and drought avoidance can be screened by a combination of leaf temperature or other rapid test of stomatal characteristics followed by carbon isotope discrimination in the most valuable materials. No single trait is adequate to improve yield in drought-prone environments, rather, a combination of characteristics is needed.     

水稻回交群体剑叶性状综合评价及QTL定位

【目的】通过对水稻剑叶性状的综合评价,明确剑叶相关性状间及与6个农艺性状的关系。检测剑叶相关性状的QTL,为优良株型品种选育,剑叶性状基因的精细定位和克隆奠定基础。【方法】以日本优质粳稻品种越光和葡萄牙粳稻地方种Bertone构建的回交群体两个世代为实验材料,利用BC₃F₁群体基因型构建遗传连锁图谱;测定亲本和BC₃F₂群体各株系剑叶SPAD、剑叶长、剑叶宽,计算剑叶长宽比、剑叶面积;利用隶属函数和标准差系数赋予权重法获得剑叶性状综合评价值(D值),分析其与6个农艺性状间的关系。分别利用单标记分析(SPA)和区间作图(IM)检测水稻剑叶相关性状QTL。【结果】在抽穗灌浆期,两亲本剑叶SPAD值呈现先升高后降低的动态变化。BC₃F₂群体的5个剑叶相关性状变异丰富,总体表现趋向轮回亲本越光。4个剑叶形态性状间相关性均达到极显著水平,与剑叶SPAD的相关性不显著。主成分和逐步线性回归分析表明剑叶宽、剑叶SPAD、剑叶长、剑叶面积是影响剑叶综合评价值(D值)的主要因子。高D值株系的株高、穗长、茎基粗和单株产量均极显著高于低D值株系,两者的分蘖数和有效穗数差异不显著。共检测到18个控制剑叶性状的QTL,分布在水稻第1、4、7和8染色体上,贡献率分布范围为4.00%~28.00%(SPA)和3.41%~27.00%(IM),除qFLSPAD1之外的17个QTL增效基因均来自Bertone。在第8染色体上的RM22720-RM404区间发现1个QTL簇,含6个主效QTL,分别为qFLL8.1、qFLL8.2、qFLA8.1、qFLA8.2、qD8.1和qD8.2。【结论】获得了剑叶宽、剑叶SPAD、剑叶长和剑叶面积4个评价剑叶性状的关键指标;明确了剑叶性状与单株产量之间的正相关关系;检测到18个剑叶相关性状QTL,位于第8染色体RM22720-RM404区间的QTL簇,是影响剑叶性状的1个重要染色体区域。     

水分胁迫对玉米叶片水分代谢的影响

试验在盆栽人工控制水分的条件下,比较了不同水分胁迫强度对不同抗旱性玉米杂交种叶片水分代谢的影响。结果表明,水分胁迫下玉米叶片水势、相对含水量、气孔导度和蒸腾速率均明显降低,随胁迫程度增强,降低幅度增大。其中叶片水势和叶片相对含水量与品种抗旱性密切相关,蒸腾速率和气孔阻力与品种抗旱性关系不明显。     

Morpho-Physiological Response of Oryza glaberrima to Gradual Soil Drying

Soil drought occurrence during dry season has been the main constraint, besides prolonged flooding during rainy season, in increasing cropping intensity and rice productivity in tropical riparian wetland. Use of drought tolerant rice genotype might be a suitable option for overcoming such problem. This study focused on the effects of gradual soil drying during early vegetative growth stage on morphological and physiological traits of five Oryza glaberrima genotypes, namely RAM12, RAM14, RAM59, RAM97 and RAM101, and two Oryza sativa subsp japonica genotypes, i.e. Koshihikari and Minamihatamochi. The plants were subjected to 6 d of gradual soil drying condition from 15 days after transplanting (DAT) to 20 DAT, and were allowed to recover until 22 DAT. Gradual soil drying reduced plant growth as indicated by dry mass accumulation. Drought reduced stomatal conductance and increased leaf rolling score of all the genotypes. All the genotypes showed comparable response on stomatal conductance, but O. glaberrima genotypes performed higher in leaf rolling recovery. Meanwhile, O. sativa genotypes decreased total leaf area and specific leaf area, but increased specific leaf weight in order to avoid further damages due to drought stress. Drought tolerance mechanisms in RAM101, RAM12, RAM59 and RAM14 were associated with leaf morpho-physiological responses, root traits and dry biomass accumulation.     

应用DNA标记分析稻飞虱的抗性基因

简要地回顾了水稻抗飞虱的遗传位点定位和作图的新进展.来自具有不同基因组的野生稻渗入系的4个抗褐飞虱基因Bph 1、 bph 2、 bph 4和Bph 9,以及4个暂定名抗褐飞虱基因Bph 10(t)、bph 11(t)、bph 12(t)和Bph 13(t),目前已被定位于水稻12条染色体中的5条.其中,Bph 1、 bph 2、 Bph 9和Bph 10(t)在水稻第12染色体的长臂上形成1个连锁区段,位于bph 2位点附近约25 cM.检测出几个对田间抗性和杀卵作用有影响的QTL.抗白背飞虱基因Wbph 1、 Wbph 2和Wbph 6(t)已经或暂时定位了.粳稻中对白背飞虱具有杀卵抗性的QTL已进行了详细的分析,在第6染色体的短臂上检测到有效的QTL,在同一位点鉴定出1个显性的杀卵基因Ovc.在杀卵基因Ovc存在时,第1染色体上的1个QTL和第5染色体上的2个QTL增加白背飞虱的卵死亡率.用DNA标记进行QTL作图可以加深人们对作物抗虫性中复杂的生理和遗传机理的理解.标记辅助选择可以加速培育具多基因抗虫性的作物,还可以将野生种中的有利抗虫特性转入改良品种中,增加作物抗虫性的持久性和遗传多样性.