Unmanaged sexual reproduction and the dynamics of genetic diversity of a vegetatively propagated crop plant, cassava (Manihot esculenta Crantz), in a traditional farming system

Occurrence of intervarietal or interspecific natural crosses has been reported for many crop plants in traditional farming systems, underlining the potential importance of this source of genetic exchange for the dynamics of genetic diversity of crop plants. In this study, we use microsatellite loci to investigate the role of volunteer seedlings (plants originating from unmanaged sexual reproduction) in the dynamics of genetic diversity of cassava (Manihot esculenta Crantz), a vegetatively propagated crop, in a traditional farming system in Guyana. A previous field study showed that farmers incorporate such plants into the germplasm for vegetative propagation, and that many of them are likely to be assigned by farmers to recognized varieties. Under strict vegetative propagation clonality of varieties is expected. The high proportion of polyclonal varieties observed suggests that incorporation of seedlings into the germplasm for propagation is a frequent event. The molecular variability assessed with microsatellite markers shows that there is high differentiation among heterozygous varieties, whereas populations of seedlings do not depart from the proportions expected under Hardy-Weinberg assumptions. Assignment of seedlings to a recognized variety on the basis of morphological similarity greatly increases genetic diversity within the variety. We argue that recombination and gene flow play a major role in the dynamics of genetic diversity of cassava in traditional farming systems. Documenting unmanaged sexual reproduction and its genetic consequences is a prerequisite for defining strategies of in situ conservation of crop plant genetic resources.     

Ecotypic differentiation reveals seed colour‐related alkaloid content in a crop wild relative

• Crop wild relatives can be a useful source of genotypes that maximise crop survival and yield in specific habitats. Lupinus angustifolius is an annual forb with crop varieties derived from a narrow genetic basis but that are cultivated worldwide. Its seeds have high nutritional value, but they naturally contain alkaloids with anti‐nutritive factors. The study of its wild populations can be useful to find genotypes that contribute to higher climate resilience and greater yield under stressing environmental conditions.
• Using a common garden, we evaluated ecotypic differentiation in four natural populations from two contrasting latitudes in terms of plant biomass, seed mass and number, alkaloid content in seeds for the three main alkaloids present in the plant and seed colour, including its possible influence on post‐dispersal predation. Correlations among traits were also assessed.
• We found differences among populations for all traits except final biomass. Northern populations had lighter seeds and a tendency to yield more seeds when they produced white seeds, compared to southern populations and variegated seeds. Regardless of latitude, populations showed differences in alkaloid concentration, with all three alkaloids found generally in high or low concentrations in each population. Proportion of white seed morphs varied in each population. Seed colour did not influence predator preference. In addition, white seed colour was related to a low alkaloid content.
• Our results evidence the existence of natural ecotypic differentiation in L. angustifolius not only due to latitudinal range, but also to local environmental factors. White seed coat colour could be used as a visual clue for identification of low‐alkaloid genotypes, a priority trait in L. angustifolius breeding programmes.

     

毛红椿天然种群有性繁殖适合度及其繁殖更新

以九连山国家自然保护区毛红椿4个天然种群为研究对象,于2006—2016年调查毛红椿有性繁殖和自然更新的情况,分析其繁殖适合度系数和个体水平适合度.结果表明:九连山毛红椿各地理种群结实单株的初始数量较少(3~9株),经70余年的繁衍发育,现存结实单株的数量仅2~10株,且来源于原始单株或子1~2代.不同种群间有性繁殖差异显著,但结实能力呈逐年下降的趋势;随群落发育成熟,土壤种子库保存与种子萌发的失利,导致能正常生长发育成熟的林木数量近乎为0.毛红椿最佳性成熟年龄为40年,早期优势明显,适合度系数早期快速增长,为2.0~2.8,但急剧减少至0.3~0.5,之后较平稳减速至近乎于0;种群间个体有性繁殖适合度(0~14株·cm2)存在较显著差异,但均较低,甚至低至0;以现有繁殖率计算,有性繁殖与更新的适合度实测值均远低于预估值.总之,受有性繁殖遗传适合度低的影响,不同种群间有性繁殖与更新能力趋同衰退;个体有性繁殖适合度进一步降低且面临更高的投资风险,现有繁殖体系因此失衡并趋于恶化.建议开展繁殖交配、授粉结实及遗传多样性评价研究,同时人为干预林分环境,于结实期清理林地枯落物,在幼苗至幼树生长过渡期适当疏伐.     

Natural hybridization between a clonally propagated crop, cassava (Manihot esculenta Crantz) and a wild relative in French Guiana

Because domestication rarely leads to speciation, domesticated populations often hybridize with wild relatives when they occur in close proximity. Little work has focused on this question in clonally propagated crops. If selection on the capacity for sexual reproduction has been relaxed, these crops would not be expected to hybridize with their wild relatives as frequently as seed-propagated crops. Cassava is one of the most important clonally propagated plants in tropical agriculture. Gene flow between cassava and wild relatives has often been postulated, but never demonstrated in nature. We studied a population of a wild Manihot sp. in French Guiana, which was recently in contact with domesticated cassava, and characterized phenotypes (10 morphological traits) and genotypes (six microsatellite loci) of individuals in a transect parallel to the direction of hypothesized gene flow. Wild and domesticated populations were strongly differentiated at microsatellite loci. We identified many hybrids forming a continuum between these two populations, and phenotypic variation was strongly correlated with the degree of hybridization as determined by molecular markers. Analysis of linkage disequilibrium and of the diversity of hybrid pedigrees showed that hybridization has gone on for at least three generations and that no strong barrier prevents admixture of the populations. Hybrids were more heterozygous than either wild or domesticated individuals, and phenotypic comparisons suggested heterosis in vegetative traits. Our results also suggest that this situation is not uncommon, at least in French Guiana, and demonstrate the need for integrated management of wild and domesticated populations even in clonally propagated crops.     

The impact of climate change on the future distribution of priority crop wild relatives in Indonesia and implications for conservation planning

The analysis of climate change impact is essential to include in conservation planning of crop wild relatives (CWR) to provide the guideline for adequate long-term protection under unpredictable future environmental conditions. These resources play an important role in sustaining the future of food security, but the evidence shows that they are threatened by climate change. The current analyses show that five taxa were predicted to have contraction of more than 30 % of their current ranges: Artocarpus sepicanus (based on RCP 4.5 in both no dispersal and unlimited dispersal scenario and RCP 8.5 in no dispersal scenario by 2050), Ficus oleifolia (RCP 4.5 5 in both no dispersal and unlimited dispersal scenario by 2080), Cocos nucifera and Dioscorea alata (RCP 8.5 in both no dispersal and unlimited dispersal scenario by 2050), and Ficus chartacea (RCP 8.5 in both no dispersal and unlimited dispersal scenario by 2050 and 2080). It shows that the climate change impact is species-specific. Representative Concentration Pathways (RCP) of greenhouse gas (GHG) emission and dispersal scenarios influence the prediction models, and the actual future distribution range of species falls in between those scenarios. Climate refugia, holdout populations, and non-analogue community assemblages were identified based on the Protected Areas (PAs) network. PAs capacity is considered an important element in implementing a conservation strategy for the priority CWR. In areas where PAs are isolated and have less possibility to build corridors to connect each other, such as in Java, unlimited dispersal scenarios are unlikely to be achieved and assisted dispersal is suggested. The holdout populations should be the priority target for the ex situ collection. Therefore, by considering the climate refugia, PAs capacity and holdout populations, the goal of keeping high genetic variations for the long-term conservation of CWR in Indonesia can be achieved.     

Failure of sexual reproduction found in micropropagated critically endangered plants prior to reintroduction: a cautionary tale

Micropropagation is a useful technique for ex situ multiplication and restoration of critically endangered plant species, but the sexual reproductive behaviour of micropropagated plants is seldom evaluated prior to reintroduction. We examined the critically endangered species Rulingia sp. ‘Trigwell Bridge’, with only three remaining plants known in the wild, as a model case to examine this issue. Abnormalities in micropropagated plants of this species related to four floral traits (lengths of sepals, petals and anthers and width of anthers). The number of pollen grains per flower of abnormal individuals was lower than in plants with apparently normal flowers (wild types), but not significantly so (P = 0.068). Pollen viability for the abnormal plant (0.87 ± 0.26%) was significantly lower than for the plants exhibiting wild‐type floral morphology (45.42 ± 4.47%). Experimental manipulations were used to examine the mating behaviour of normal and abnormal plants. The results showed that both male and female reproductive failure was linked to individuals exhibiting abnormal flowering attributes. Such aberrant reproductive performance in a micropropagated rare species predicates caution when using micropropagated plants in reintroduction programmes, highlighting the importance of screening for reproductive normality prior to release of micropropagated plants (especially for critically endangered species where reliance on in vitro propagation methods is often a necessity). © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 278–284.     

The conservation of redundancy in genetic systems: effects of sexual and asexual reproduction

The relationship between probability of survival and the number of deleterious mutations in the genome is investigated using three different models of highly redundant systems that interact with a threatening environment. Model one is a system that counters a potentially lethal infection; it has multiple identical components that act in sequence and in parallel. Model two has many different overlapping components that provide three-fold coverage of a large number of vital functions. The third model is based on statistical decision theory: an ideal detector, following an optimum decision strategy, makes crucial decisions in an uncertain world. The probability of a fatal error is reduced by a redundant sampling system, but the chance of error rises as the system is impaired by deleterious mutations. In all three cases the survival profile shows a synergistic pattern in that the probability of survival falls slowly and then more rapidly. This is different than the multiplicative or independent survival profile that is often used in mathematical models. It is suggested that a synergistic profile is a property of redundant systems. Model one is then used to study the conservation of redundancy during sexual and asexual reproduction. A unicellular haploid organism reproducing asexually retains redundancy when the mutation rate is very low (0001 per cell division), but tends to lose high levels of redundancy if the mutation rate is increased (001 to 01 per cell division). If a similar unicellular haploid organism has a sexual phase then redundancy is retained for mutation rates between 0001 and 01 per cell division. The sexual organism outgrows the asexual organism when the above mutation rates apply. If they compete for finite resources the asexual organism will be extinguished. Variants of the sexual organism with increased redundancy will outgrow those with lower levels of redundancy and the sexual process facilitates the evolution of more complex forms. There is a limit to the extent that complexity can be increased by increasing the size of the genome and in asexual organisms this leads to progressive accumulation of mutations with loss of redundancy and eventual extinction. If complexity is increased by using genes in new combinations, the asexual form can reach a stable equilibrium, although it is associated with some loss of redundancy. The sexual form, by comparison, can survive, with retention of redundancy, even if the mutation rate is above one per generation. The conservation and evolution of redundancy, which is essential for complexity, depends on the sexual process of reproduction.     

Fine scale genetic structure in the wild ancestor of maize (Zea mays ssp. parviglumis)

Analysis of fine scale genetic structure in continuous populations of outcrossing plant species has traditionally been limited by the availability of sufficient markers. We used a set of 468 SNPs to characterize fine‐scale genetic structure within and between two dense stands of the wild ancestor of maize, teosinte (Zea mays ssp. parviglumis). Our analyses confirmed that teosinte is highly outcrossing and showed little population structure over short distances. We found that the two populations were clearly genetically differentiated, although the actual level of differentiation was low. Spatial autocorrelation of relatedness was observed within both sites but was somewhat stronger in one of the populations. Using principal component analysis, we found evidence for significant local differentiation in the population with stronger spatial autocorrelation. This differentiation was associated with pronounced shifts in the first two principal components along the field. These shifts corresponded to changes in allele frequencies, potentially due to local topographical features. There was little evidence for selection at individual loci as a contributing factor to differentiation. Our results demonstrate that significant local differentiation may, but need not, co‐occur with spatial autocorrelation of relatedness. The present study represents one of the most detailed analyses of local genetic structure to date and provides a benchmark for future studies dealing with fine scale patterns of genetic diversity in natural plant populations.     

Barriers to sexual reproduction in Polygonum viviparum: a comparative developmental analysis of P. viviparum and P. bistortoides

Polygonum viviparum is widely distributed in arctic and alpine regions of the northern hemisphere. Fruit set has never been observed in North American populations and has been reported only very rarely in Europe. Although this species is extremely well studied, the impediments to successful fruit production are unknown. We investigated the sexual reproductive process in P. viviparum growing in the southern Colorado Rocky Mountains. For comparison, we also examined this process in the sympatric congener P. bistortoides, in which reproduction is exclusively sexual. Lack of viable fruit production in P. viviparum has no single developmental explanation; defects occur in each of the processes and structures associated with sexual reproduction studied, yet, these processes and structures also appear to function normally in at least some flowers or individuals. Development is abnormal in many ovules of P. viviparum, however, comparison with P. bistortoides shows that these abnormalities do not contribute to differences in seed production between the two species. The virtual absence of sexual reproduction in P. viviparum appears to be due largely to a low rate of fertilization and to embryo/fruit abortion.     

The importance of sexual and asexual reproduction in the recent evolution of Allium vineale

In the weedy plant species Allium vineale (wild garlic), individuals may simultaneously produce sexually and asexually derived offspring, by seed and bulbils, respectively. In this study, genetic and genotypic diversity was determined in samples from 14 European A. vineale populations using nuclear (RAPD) and cytoplasmic (PCR-RFLP of cpDNA) markers to investigate the importance of the different reproductive modes. In the whole sample, 77 nuclear multilocus genotypes and four chloroplast haplotypes (chlorotypes) were found. Populations exhibited a high degree of subdivision for nuclear and cytoplasmic markers as estimated from hierarchical F-statistics; at the same time, identical chlorotypes could be found in populations separated by large distances. Genotypic diversity was significantly lower than expected under free recombination in almost all populations, indicating that recruitment into populations is mostly by asexually produced offspring. Nevertheless, within each chlorotype, the distribution of markers from pairs of nuclear loci was incompatible with a purely clonal structure, suggesting that many multilocus genotypes have originated by sexual recombination rather than by mutation within asexual lineages. It is argued that the weedy habit of A. vineale is likely to have favored bulbil reproduction, whereas sexually generated genotypes may have facilitated local adaptation during the species' expansion across Europe.     

The relative contributions of sexual reproduction and clonal propagation in Opuntia rastrera from two habitats in the Chihuahuan Desert

1 The clonal cactus Opuntia rastrera shows predominantly sexual reproduction in grasslands (GH) and clonal propagation in nopaleras (NH). We assessed the effects of light, herbivory, water availability and the habitat an offspring came from on the survival and growth of sexual or clonal offspring (i.e. seedlings and cladodes), through 3- and 4-year common garden and short-term greenhouse experiments.
2 Shading by nurse plants increased seedling survival in the field by an order of magnitude, and a small additional advantage due to predator protection by grasses was observed. Strong herbivory transforms a facultative nurse–protégé relationship for seedlings into an obligatory one.
3 In the greenhouse seedlings grew better under shade, but in the field the production of the first cladode was delayed in seedlings in the more shaded GH. Competition for soil resources may be more intense under a dense grass tussock than under a open shrub, thus affecting the nurse–protégé relationship. Seedling survival under nurse plants was similar in GH and NH, but higher plant cover suggests that a larger number of seedlings will establish in GH in the long term.
4 Cladode survival was higher in NH. Cladodes were more successful than seedlings at establishing in intercanopy areas, possibly due to physiological differences as well as their ability to survive partial predation. Cladode survival in intercanopy areas may explain the enhanced clonal propagation in the more open NH scrubland, together with their susceptibility to the flooding which affects GH.
5 The high seedling and cladode survival in the greenhouse experiments contrasted with that observed in the field, indicating that survival is determined by the interaction between herbivores, plants and abiotic conditions rather than the physiological aptitude of the plants.     

The role of sexual and clonal reproduction in maintaining population in Fritillaria camtschatcensis (L.) Ker-Gawl. (Liliaceae)

Fritillaria camtschatcensis can reproduce by means of both sexual reproduction and clonal multiplication. Despite prolific seed production, no seedlings have been found for several years in natural populations on Mt Hakusan. The purposes of this paper are to clarify: (i) whether population size is maintained mainly by clonal multiplication; and (ii) if this is the case, to what extent occasional seedling establishment affects population growth rate and population structure. Two permanent quadrats were placed in subalpine meadows in 1992 on Mt Hakusan. Plant size, location and reproductive states for all ramets in the quadrats were recorded every year. Projection matrices were created based on field census, and computer simulation experiments were performed. Fritillaria camtschatcensis had two types of flower, male flower and cosexual flower, and they were changeable. This is the first report on sex lability in Fritillaria. Clonal growth was more closely correlated with life-history stages, especially with sexual states than with plant size. The population growth rate, , was 1.006 for the Mizuyajiri population and 1.047 for the Nanryu population, respectively. Seedlings were found in 1996 for the first time. These facts indicate that populations of F. camtschatcensis on Mt Hakusan can usually be maintained by clonal multiplication. However, it is not yet certain whether seeds germinate every year or whether a flush of seedling emergence occurs once in every few years in natural populations. Computer simulation revealed that: (i) there was a critical germination rate above which population growth rate suddenly increased; and (ii) occasional seedling establishment could provide almost the same contribution to population growth rate as that of annual seedling establishment. These results suggest that population size can be maintained mainly by clonal multiplication, and the role of sexual reproduction lies beyond maintaining the population size in F. camtschatcensis.     

Cadmium,copper and zinc toxicity effects on growth,proline content and genetic stability of Solanum nigrum L., a crop wild relative for tomato; comparative study

Plants like other organisms are affected by environmental factors. Cadmium, copper and zinc are considered the most important types of pollutants in the environment. In this study, a comparison of growth and biochemical parameters between the crop wild relative (CWR) Solanum nigrum versus its cultivated relative Solanum lycopersicum to different levels of Cu, Zn and Cd stress were investigated. The presence of ZnSO₄ and CuSO₄ in Murashige and Skoog medium affected severely many growth parameters (shoot length, number of roots and leaves, and fresh weight) of both S. nigrum and S. lycopersicum at high levels. On the other hand, CdCl₂ significantly reduced most of the studied growth parameters for both species. S. nigrum exhibited higher tolerance than S. lycopersicum for all types of stress. In addition, results show that as stress level increased in the growing medium, proline content of both S. nigrum and S. lycopersicum increased. A significant difference was observed between the two species in proline accumulation as a result of stress. In addition, a higher accumulation rate was observed in the crop wild relative (S. nigrum) than in cultivated S. lycopersicum. Changes in Inter-simple sequence repeat (ISSR) pattern of CuSO₄ treated S. nigrum and S. lycopersicum plants were also observed. In conclusion, based on growth and biochemical analysis, S. nigrum showed higher level of metals tolerance than S. lycopersicum which indicates the possibility of using it as a crop wild relative for S. lycopersicum.     

浙江丽水中国石龙子的食性、两性异形和雌性繁殖

丽水分布的中国石龙子（Ｅｕｍｅｃｅｓ ｃｈｉｎｅｎｓｉｓ）摄入的食物均为无脊椎动物,分别属于环节、软体和节肢动物,涉及３０余科,成体和幼体的食物生态位宽度分别为７．２６和６．６９,成体的幼体的食物生态们重叠度为０．５９。性成熟雄性个体大于雌体。成雄和幼体的头长和头宽随体长ＳＶＬ的增长速率大于成雌,成雄头长随ＳＶＬ的增长速度显著大于幼体,成雌和幼体的头长随ＳＶＬ的增长速率无显著差异。成雄头部大于成雌     

岷江干旱河谷多苞蔷薇生长与繁殖特征

通过大面积野外踏查和样方调查,分析了岷江干旱河谷乡土灌木多苞蔷薇种群的分布、生长繁殖特征及其空间差异性.结果表明,多苞蔷薇广泛分布于岷江干旱河谷汶川绵褫至松潘镇江关约170 km长、海拔1 200～2 600 m的空间范围内,其株高、基径、年龄、冠幅、萌蘖株数和结实量的平均值分别为(123.33±3.13) cm、(12.49±0.40) mm、(3.34±0.16) a、(1.30±0.13) m²、(5.07±0.39)株/丛和(171.47±25.90)果/丛.随着海拔高度和纬度增加,其株高、基径、冠幅和结实量增加,而萌蘖株数没有显著变化.多苞蔷薇有性结实和无性萌蘖能力随环境空间变化的不匹配性与其生长密切相关,可能与多变环境下资源分配利用的平衡机制有关;以丛进行年龄统计,多苞蔷薇缺少幼龄个体,种群发展并不稳定,但随海拔高度升高,萌蘖株年龄结构均呈“金字塔”型,且较大龄级的萌株比例略增,显示出良好的种群持续性.综合分析发现,多苞蔷薇生长与繁殖的空间差异性是对干旱河谷长期适应的结果,既为种群持续提供了保证,也为种群扩大奠定了基础.     

Auxosporulation of Licmophora communis (Bacillariophyta) and a review of mating systems and sexual reproduction in araphid pennate diatoms

The auxosporulation of Licmophora communis is allogamous and dioecious. Pairing between sessile, shortstalked cells of compatible clones is followed by meiosis and gametogenesis, to form two gametes in each gametangium. The behavior of the gametes differs between the gametangia. In the male gametangium, the gametes detach from the frustule, round up, and migrate out of the gametangium after its dehiscence at the broader, unattached pole. In the female gametangium, both gametes remain attached to the adjacent theca over almost their whole length and do not move. Plasmogamy therefore occurs within the female gametangium and this is where the zygotes are formed and remain. After fertilization, the zygotes detach from the thecae of the female gametangia, contract, and become ellipsoidal, before expanding parallel to the apical axis of the gametangium. We review the types of auxosporulation in other pennate diatoms and the systems used for classifying these. Dioecy and cis‐type anisogamy (in which one gametangium produces active gametes and the other produces passive gametes), as in L. communis, are probably primitive within the pennate group (although there is no information on the AsterionellopsisRhaphoneis clade). However, size can also be restored in various araphid pennates by allogamous sexual reproduction involving the formation of only one gamete per gametangium, or in rare cases by automixis or (apparently) vegetative enlargement.     

Genome-wide association mapping identifies the genetic basis of discrete and quantitative variation in sexual weaponry in a wild sheep population

Understanding the genetic architecture of phenotypic variation in natural populations is a fundamental goal of evolutionary genetics. Wild Soay sheep (Ovis aries) have an inherited polymorphism for horn morphology in both sexes, controlled by a single autosomal locus, Horns. The majority of males have large normal horns, but a small number have vestigial, deformed horns, known as scurs; females have either normal horns, scurs or no horns (polled). Given that scurred males and polled females have reduced fitness within each sex, it is counterintuitive that the polymorphism persists within the population. Therefore, identifying the genetic basis of horn type will provide a vital foundation for understanding why the different morphs are maintained in the face of natural selection. We conducted a genome-wide association study using ～36000 single nucleotide polymorphisms (SNPs) and determined the main candidate for Horns as RXFP2, an autosomal gene with a known involvement in determining primary sex characters in humans and mice. Evidence from additional SNPs in and around RXFP2 supports a new model of horn-type inheritance in Soay sheep, and for the first time, sheep with the same horn phenotype but different underlying genotypes can be identified. In addition, RXFP2 was shown to be an additive quantitative trait locus (QTL) for horn size in normal-horned males, accounting for up to 76% of additive genetic variation in this trait. This finding contrasts markedly from genome-wide association studies of quantitative traits in humans and some model species, where it is often observed that mapped loci only explain a modest proportion of the overall genetic variation.     

Reproductive ageing and sexual selection on male body size in a wild population of antler flies (Protopiophila litigata)

Little is known about the importance of trade-offs between ageing and other life history traits, or the effects of ageing on sexual selection, particularly in wild populations suffering high extrinsic mortality rates. Life history theory suggests that trade-offs between reproduction and somatic maintenance may constrain individuals with higher initial reproductive rates to deteriorate more rapidly, resulting in reduced sexual selection strength. However, this trade-off may be masked by increased condition dependence of reproductive effort in older individuals. We tested for this trade-off in males in a wild population of antler flies (Protopiophila litigata). High mating rate was associated with reduced longevity, as a result of increased short-term mortality risk or accelerated ageing in traits affecting viability. In contrast, large body size was associated with accelerated ageing in traits affecting mating success, resulting in reduced sexual selection for large body size. Thus, ageing can affect sexual selection and evolution in wild populations.