Climatic adaptation of trees: rediscovering provenance tests

Summary Common garden testing of populations of different origin started with forest trees more than two hundred years ago. Since then, so-called provenance tests have been established with most commercially important species. Beyond the strictly silvicultural goals, the tests offer excellent opportunities to study intraspecific genetic variation patterns and represent probably the most powerful available tool for testing hypotheses of climatic adaptation in trees.Analysis of adaptive traits (mostly juvenile height growth) in provenance experiments indicate the existence of very effective constraints on adaptedness. The performance of populations plotted against an ecological-climatic factor exhibits a characteristic pattern and can be described by response functions. The population average of a fitness-related trait for a locally adapted population is often significantly lower than that of populations from other environments; usually the ones from milder climate perform better. The phenomenon is interpreted as adaptation lag. Suboptimal adaptation is compensated by a high level of genetic diversity. Molecular genetic studies confirm the high level of allelic and individual genetic diversity in forest trees. A consequence of individual homeostasis, phenotypic stability of populations is usually also high; the sensitivity to environmental changes is generally moderate. Phenotypically stable populations are valuable not only because of a wider range of potential cultivation but specifically because of a greater ability to adjust to unexpected changes. This trait should receive more attention in the future for obvious reasons.The maintenance of a high within-population genetic variance is favored by the genetic system of the investigated species (effective gene flow, outbreeding, high genetic load, etc.). Random events and long-lasting biotic interactions are further effects impairing the efficiency of natural selection.In view of expected climate instability, genetic adaptability of forest trees causes serious concern due to their long lifespan compared to the rapidity of expected changes in environmental conditions. The potential of provenance tests to interpret long-term adaptational processes should be utilized to analyze, model and predict response of trees to climate change. Although seldomly appreciated, provenance research might be among the most important contributions of forestry to biological sciences.     

Complex constitutive nature of Japanese upland rice, Oryza sativa L.

Two rice ecotypes, the so-called lowland and upland populations, which carry different isozyme genotypes mostly at a single locus, are cultivated in Japan. The aim of this study was to examine the origin and the mechanism for keeping these genetic differences. The upland population is cultivated in upland fields and carries a different allele for a particular isozyme gene, Pgd-1, which has never been found in the lowland population. RFLP markers showed a weak trend for genetic differentiation between the two ecotypes. On the other hand, morphological, and physiological traits showed marked differences between the two ecotypes. Furthermore, based on the genotypic difference, two Japonica subgroups are defined in the upland population. Subgroup I is the minor group and carries key lowland characters, including the genotype for PGD. Subgroup II carries different traits and the genotype for PGD of the alternative subgroup. As an allelic difference for Pgd-1 is known to occur between the two ecospecies, Tropical (Tr) and Temperate (Tm) Japonicas, upland cultivars can be classified by diagnostic characters which distinguish a variety into Tr or Tm type. The upland population consists of three types of cultivars, Tr-, Tm- and intermediate-type. In contrast, the lowland population consists of a uniform Tm type Japonicas. As Japanese upland cultivars still have an isozyme allele specific to the Tr type, the upland population has a rather complex constitution which is presumably now being introgressed by lowland genetic material, but still represents a major difference at some genetic levels. Upland rice carries several stress-resistant genes which would be useful for lowland rice breeding. The genetic difference would be efficient for tagging upland specific traits by upland specific genetic markers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolution and adaptedness in a facultatively apomictic grass, Poa pratensis L.

Summary Facultative apomixis confers great adaptedness and evolutive potential on biotypes belonging to the Poa pratensis L. agamic complex. Pioneering and recent research have dealt with and sometimes elucidated different aspects of apomictic expression, such as cytological mechanisms, distribution patterns, genetic control and environmental effects, in this species. We carried out an investigation on the mode of reproduction expressed in Italian natural populations, cultivars and selected genotypes together with an extensive time (56 years) and space (six countries) review in order to obtain a comprehensive picture of apomixis expression in P. pratensis. Histograms of the estimations were prepared. They showed that, due to breeder and natural selection, both cultivars and natural populations very frequently expressed a high degree of apomixis. Variations in the degree of apomixis across generations, different pollen donors, environments and estimation levels were generally low. It would, therefore, seem that the tenet which holds that apomixis in P. pratensis is strongly influenced by external factors should be reconsidered. Despite the fact that little is known of the genetic, physiological and molecular control underlying apomixis, the overall picture that emerges from our study helps to explain how the balance between sexual and asexual reproduction confers extreme versatility, adaptedness and evolutive capacity on this remarkable grass.     

Challenges and opportunities for conservation of forest genetic resources

Increased use of forest resources and a shrinking forestland base threaten the sustainability of forest genetic resources and highlight the importance of conservation and sustainable management of these resources. As forest trees are normally the keystone species of forest ecosystems, their continued existence is essential for many floral and faunal associations of these ecosystems. Major concepts, challenges and opportunities for conservation of forest genetic resources are briefly discussed in this paper. The major challenges include population decline and population structure changes due to forest removal and conversion of forest land to other uses,forest fragmentation, forestry practices, climate change, disease conditions,introduced pests, atmospheric pollution, and introgressive hybridization. Developing scientifically sound conservation strategies, maintaining minimum viable population sizes, and deployment of genetically engineered organisms represent other important challenges in conservation. The usefulness of various biochemical and molecular genetic markers, adaptive traits, and genetic diversity measures for developing conservation strategies for in situ and ex situ genetic resource conservation are also discussed. Major opportunities for conservation of forest genetic resources include: use of molecular genetic markers and adaptive traits for developing conservation strategies; in situ conservation through natural reserves,protected areas, and sustainable forest management practices; ex situ conservation through germplasm banks, common garden archives, seed banks, DNA banks, and tissue culture and cryopreservation; incorporation of disease, pest, and stress tolerance traits through genetic transformation;plantation forestry; and ecological restoration of rare or declining tree species and populations. Forest genetic resource conservation and resource use should be considered complementary rather than contradictory to each other. This revised version was published online in July 2006 with corrections to the Cover Date.     

Recurrent selection for broad adaptation affects stability of oat

A recurrent selection program for adaptation to diverse environments was successful in improving mean oat (Avena sativa L.) grain yield within and across testing environments. The objectives of this research were to determine if this selection program also resulted in changes in other agronomic traits or altered yield stability. Additionally, we investigated how selection modified the response of genotypes to climatic conditions. We evaluated random samples of 100 families from the original population and each of three selection cycle populations in replicated yield trials in Idaho, Iowa, and Norway for two years. Yield stability was assessed via joint regression analysis and superiority analysis. For each cycle, genetic relationships among yields observed indifferent environments were assessed by estimating phenotypic correlations between pairs of target environments. The effect of climate variables on genotype-by-environment interaction (GEI) responses was determined with partial least squares regression. Selection resulted in a small increase in mean heading date, a decrease in mean test weight, and no change in total above-ground biomass or plant height. Genotypic regression coefficients on environmental indices and deviations from regression were larger in the last cycle population, but superiority analysis demonstrated that selection significantly improved the adaptability of the population to the target testing environments. Improved adaptation was also demonstrated by increased phenotypic correlations among the most divergent pairs of environments in the later cycles. Partial least squares regression of GEI effects on climate variables suggested that later cycle families tended to respond more favorably to cooler than average conditions than the original population. Selection resulted in improved yield stability as well as improved mean yield. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic basis of the evolution of adaptedness in plants

Summary Adaptedness is both complexly inherited and much affected by environment: consequently the genetic mechanisms that have led to improvements in adaptedness have been difficult to identify and to quantify. Recently it has been shown that marker assisted dissection of adaptedness based on changes in the frequencies of discretely inherited alleles of loci of various kinds (e.g. allozyme, restriction fragment, microsatellite loci) is practicable. I will illustrate marker assisted analysis of the genetic basis of adaptedness with a sample of allozyme data from three species groups, two heavily selfing groups (two wild Avena species and barley) and one outcrossing species (corn, maize). The results lead to three main conclusions: (1) that the single most important genetic mechanism in all three species groups was the assembly of favorable epistatic combinations of alleles of different loci by means of recurring cycles of selection, intercrossing superior selects, and inbreeding to near homozygosity leading to stable superior multilocus genotypes adapted to specific habitats; (2) that exploitation of favorable interactions among alleles of the same locus played a significant role in tetraploid A. barbata and probably also in single-cross maize hybrids; (3) that purifying selection (elimination of deleterious alleles) played a small role in all three species groups. These results indicate that marker alleles provide applied breeders with effective ways to identify, track, and incorporate regions of chromosomes with favorable effects of adaptedness into improved cultivars.     

Molecular markers to exploit genotype–environment interactions of relevance in organic growing systems

One of the substantial differences between conventional and organic growing systems is the degree to which the farmer can control biotic and abiotic stresses; for organic growing systems varieties are needed with a broad adaptation to annually varying factors, while at the same time a good specific adaptation is necessary with respect to more constant climate and soil conditions. This combination of requirements implies that varieties for organic farming need to be better characterised with respect to genotype × environment interactions than varieties for conventional farming. Such interactions, which often are found for quantitatively expressed traits, are in general difficult to deal with in phenotypic selection. New approaches for QTL analyses (e.g. using physiological models) facilitate estimation of effects of genes on a trait (the phenotype) as a response to environmental influences. From such analyses, markers can be identified which may help to predict the trait expression of a plant genotype in relation to defined environmental factors. The application of markers to select for loci with specific interactions with the environment could, therefore, be especially important for plant breeders targeting organic farming systems.     

Analyses of genetic diversity in Cuban rice varieties using isozyme, RAPD and AFLP markers

A survey of the genetic diversity among the major cuban rice cultivars was conducted using isozyme, RAPD and AFLP markers. Polymorphisms were detected for esterases, peroxidases, alcohol dehydrogenases and polyphenoloxidases systems; 21 RAPD primers and four AFLP primer combinations. Heterozygosity arithmetic mean value (H_av(p)), the effective multiplex ratio (EMR) and the marker index (MI), were calculated for isozyme, RAPD and AFLP markers. The mean value of genetic similarity among the different varieties was 0.92 for isozyme, 0.73 for RAPD and 0.58 for AFLP analyses. Thus, AFLP were able to detect polymorphisms with higher efficiency than RAPD (+15%) and isozyme (+34%). Data from the isozyme, RAPD and AFLP analyses were used to compute matrices of genetic similarities. The efficiency of the UPGMA for the estimation of genetic relatedness among varieties was supported by cophenetic correlation coefficients. The resulting values indicated that the distortion level for the estimated similarities was minimal. The correlation coefficients obtained by the Mantel matrix correspondence test, which was used to compare the cophenetic matrices for the different markers, showed that estimated values of genetic relationship given for isozyme and RAPD markers (r = 0.89), as well as for AFLP and RAPD markers (r = 0.82) were properly related. However, AFLP and isozyme data showed only moderate correlation (r = 0.63). Although the genetic variability found among the different cultivars was low, both RAPD and AFLP markers proved to be efficient tools in assessing the genetic diversity of rice genotypes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evaluating genetic variation and relationships among two bromegrass species and their hybrid using RAPD and AFLP markers

The two most widely grown bromegrass species in North America are smooth bromegrass (Bromus inermis Leyss.), a hay type grass, and meadow bromegrass (Bromus riparius Rehmann), a pasture type grass. Recently a hybrid bromegrass population between the two species has been produced as a dual-purpose hay-pasture grass. Molecular markers have the potential to improve selection procedures to enhance bromegrass breeding. The objective of this study was to use RAPD and AFLP markers to determine genetic relationships and variations among bromegrass populations. Forty-three RAPD markers from 21 primers and 83 AFLP markers from seven primer combinations were used. Both marker types were able to group the individuals into their respective populations. The relationships among the individuals within each of the populations were not similar between the two marker types. Analysis of molecular variance (AMOVA) detected greater within-population variation than among-population variation for both marker types. The highest variation was observed in the hybrid population followed by meadow and then smooth bromegrass. The inter-population distance from both markers indicated that the highest genetic distance was between meadow and smooth bromegrass and lowest between smooth and hybrid bromegrass, which reflect the breeding history of the hybrid population. This study showed that both markers are capable of differentiating bromegrass genotypes into their respective populations, detecting genetic variation and relationships of the populations. Results of this study suggest that these two markers can be used in the future to enhance the current breeding practices in bromegrass, however, AFLP markers would be the marker of choice due to the high number of polymorphic markers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Isozyme variation in Helianthus petiolaris and sunflower,H. annuus

Summary Helianthus petiolaris is a wild species used in genetic improvement of sunflower, as a donor of cytoplasmic male sterility and of genetic resistance to diseases. Isozyme variation for ADH, ACP, EST, GDH, LAP, PGI, PGD and SKDH in this species was studied using starch gel electrophoresis. The patterns thus obtained were compared with zymograms of inbred lines, hybrids and open pollinated varieties of H. annuus. The same alleles for EST and SKDH isozymes were found in both species, while ACP showed an allele that has not been found in sunflower. The rest of the isozyme systems showed both common alleles and characteristic ones for each species. ACP, GDH and PGD were monomorphic in H. petiolaris, while ADH and LAP were monomorphic in H. annuus. The isozyme markers obtained here could be useful in breeding programs involving interspecific crosses, and studies on introgression and on genetic variation in other populations of this wild species.     

Breeding Strategies for Adaptation of Pearl Millet and Sorghum to Climate Variability and Change in West Africa

Semi‐arid and subhumid West Africa is characterized by high inter‐annual rainfall variability, with variable onset of the rainy season, somewhat more predictable endings, and drought or excess water occurrence at any time during the growing season. Climate change is predicted to increase this variability. This article summarizes options for plant breeders to enhance the adaptation of pearl millet (Pennisetum glaucum [L.] R. Br.) and sorghum (Sorghum bicolor [L.] Moench) to climate variability in West Africa. Developing variety types with high degrees of heterozygosity and genetic heterogeneity for adaptation traits helps achieving better individual and population buffering capacity. Traits that potentially enhance adaptive phenotypic plasticity or yield stability in variable climates include photoperiod‐sensitive flowering, plastic tillering, flooding tolerance, seedling heat tolerance and phosphorus efficiency. Farmer‐participatory dynamic gene pool management using broad‐based populations and diverse selection environments is useful to develop new diverse germplasm adapted to specific production constraints including climate variability. For sustainable productivity increase, improved cultivars should respond to farmer‐adoptable soil fertility management and water harvesting techniques. Larger‐scale, on‐farm participatory testing will enable assessments of varietal performance under evolving climatic variability, provide perspective on needs and opportunities and enhance adoption. Strengthening seed systems will be required to achieve sustainable impacts.     

Farmers' perceptions of and responses to environmental change in southwest coastal Bangladesh

Coastal Bangladesh is highly vulnerable to climate change and salinisation; hence, farm‐level adaptation is critically important. Farmers' perceptions of and responses to environmental change were investigated in two villages in Khulna District. Perceived environmental trends included increases in temperature, extreme weather events, soil toxicity, erratic rainfall and scarcity of water for irrigation. Perceptions of climate trends were consistent with measured trends in Khulna. On‐farm adaptation strategies included adjusting planting dates, excavating trenches in rice‐fields, adopting new crops, salinity‐reducing technologies, livestock‐rearing and home‐yard cropping. Non‐farm adaptation strategies included wage employment, short‐term migration and self‐employment. Adaptation was facilitated by income‐earning opportunities, training, and credit, and impeded by lack of access to water, markets, capital, and extension services. Farmers suggested policy support for dissemination of stress‐tolerant cultivars, access to irrigation, and price stabilisation or crop insurance to assist adaptation. While the study shows an impressive degree of awareness and adaptation, external support is needed to increase adaptive capacity.     

苦荞SSR分子遗传图谱的构建及分析

构建苦荞遗传连锁图谱,为今后有关苦荞基因组结构、重要农艺性状QTL定位、分子标记辅助育种和基因克隆等研究工作奠定基础。以栽培苦荞‘滇宁一号’和苦荞野生近缘种杂交产生的119份F4代分离材料为作图群体,利用SSR分子标记来构建苦荞的分子遗传连锁图谱。本研究构建的连锁图谱包含15个连锁群,由89个标记组成,其中偏分离的标记有22个,占24.7%,每条连锁群上的标记在2~16之间。连锁群长度在6.9~165.8 cM的范围,覆盖基因组860.2 cM,总平均长度9.7 cM。本研究构建了首张苦荞SSR遗传连锁图谱,为苦荞QTL定位、基因克隆、遗传选育等研究奠定了基础。     

Breeding winter hardy grasses

Summary Perennial grasses are vital for Norwegian agricultural production. The nature and extent of winter damage on grasslands is highly dependent on climatic conditions, and determines both persistency and yield. Physical stresses such as frost and ice encasement predominate in coastal regions with an unstable winter climate, while biotic stresses such as low temperature fungi are more common in the inland regions. Development of hardening depends on plant adaptation and climatic conditions during autumn and winter. New winter-hardy cultivars should be bred for wide adaptation to winter stresses. The genetic background for the most important character, freezing tolerance, seems to be of polygenic nature with mainly additive gene action. Selection for increased freezing tolerance has been effective over generations in grasses, and in most grass species ample variation still exists to be exploited by breeding. However, in some species like perennial ryegrass, modern biotechnological methods should be used to improve freezing tolerance and winter hardiness.     

Construction of a linkage map for watermelon (Citrullus lanatus (Thunb.) Matsum & Nakai) using random amplified polymorphic DNA (RAPD)

Summary A linkage map for watermelon (Citrullus lanatus) was constructed on the basis of RADP, ribosomal DNA restriction fragment length polymorphism (RFLP), isozyme, and morphological markers using F₁BC1. A segregating population of 78 individuals was the result of a backcross of a cultivated inbred line (H-7; Citrullus lanatus; 2n=22) and a wild form (SA-1; C. lanatus; 2n=22), in which the latter was the recurrent (male) parent. A total of 69 RAPD, one RFLP, one isozyme, and three morphological markers was found to segregate in the BC1 population. Linkage analysis revealed that 62 loci could be mapped to 11 linkage groups that extended more than 524 centimorgans (cM), while 12 loci segregated independently of all other markers. The locus for exocarp color was linked to two RAPD markers within a region of 5 cM on linkage group 4. The locus for flesh color was linked to a RAPD marker within a region of 30 cM on linkage group 6. The isozyme marker GOT was located on the linkage group 1. Linkage group 2 contained a locus for ribosomal DNA within 5 cM of a RAPD marker. Half of the RAPD markers on the linkage group 7 displayed severely distorted segregation. The construction of linkage map using molecular markers is necessary for the breeding of watermelon to introduce useful gene of wild watermelon efficiently. However the linkage map that was constructed for the most part on the basis of RAPD markers could not cover significant parts of the genome, the linkage map provides breeders of watermelons the possibility of tagging useful agronomic traits, as well as the gene for exocarp color.Abbreviations RAPD random amplified polymorphic DNA - RFLP restriction fragment length polymorphism - GOT glutamate oxaloacetate transaminase - MDH malate dehydrogenase - ACP acid phosphatase - 6PGH 6-phosphogluconate dehydrogenase     

不同寄主致病疫霉菌群体遗传结构的SSR比较分析

用SSR分子标记技术对采集自中国不同地区不同寄主来源的60个致病疫霉菌(Phytophthora infestans)群体遗传多样性进行比较分析。结果显示,番茄致病疫霉菌群体Nei氏基因多样性指数、Shannon信息指数及有效等位基因数等遗传多样性指数(分别为0.1528、0.2592、1.2170)均高于马铃薯致病疫霉菌（分别为0.1395、0.2447、1.1913）,表明番茄致病疫霉菌群体遗传多样性高于马铃薯致病疫霉菌群体;聚类分析结果表明：SSR标记分析的遗传系谱与菌株对甲霜灵敏感性、交配型无相关性,而与MtDNA单倍型有一定的相关性。     

Estimating the genetic relationship of hybrid bromegrass to smooth bromegrass and meadow bromegrass using RAPD markers

The two most widely grown bromegrass species in North America are smooth bromegrass, a hay type grass, and meadow bromegrass, a pasture type grass. Hybrid populations between these two species have been developed through hybridization and recurrent selection. The objective of this study was to determine the genetic relatedness of the hybrid bromegrass population S‐9073M to its parental populations using random amplified polymorphic DNA (RAPD) markers and to determine genetic variation within and between populations. Individuals from each of the three populations were genotyped at 43 polymorphic RAPD loci. One of the RAPD fragments was meadow bromegrass‐specific. Cluster analysis showed three groups representing the two parental populations and the hybrid population. An analysis of molecular variance (AMOVA), showed that the hybrid population had the highest within‐population variation, followed by meadow bromegrass and smooth bromegrass. The interpopulation genetic distance (phi‐statistic =Φ_st) was highest between meadow and smooth bromegrass and lowest between smooth and hybrid bromegrass. The hybrid population was genetically intermediate to smooth bromegrass and meadow bromegrass, but closer to smooth bromegrass, possibly reflecting the selection criteria used in the development of this hybrid.     

植物高温胁迫响应分子机制研究

作为固着生物,植物需要面对持续变化的环境,不断地受到温度和其他非生物因素的影响。未来植物所面临的平均环境温度将会随着全球气候变暖逐渐上升,因此高温愈发成为影响植物生长发育、地理分布及产量的非生物胁迫因子。了解植物对高温胁迫的适应机制,有助于开发耐热品种,改善温暖气候地区植物的生长状况以及提高作物的生产力。植物体中,在分子水平,转录组学及蛋白质组学联合揭示了mRNA以及蛋白水平的植物应答高温胁迫的基因表达模式变化;另外高温胁迫广泛涉及植物体内多种信号转导机制。因此,本综述主要从植物响应高温胁迫分子水平上的转录组、蛋白质组学方面,以及信号转导等方面全面总结了植物响应高温胁迫的研究进展,并对未来的研究前景进行了展望,为了解近几年高温胁迫研究动态和高温胁迫常见研究方式,解析植物高温胁迫耐受机理以及耐高温品种的培育提供参考。     

Genetic characterization and gene flow in different geographical-distance neighbouring natural populations of wild soybean (Glycinesoja Sieb. & Zucc.) and implications for protection from GM soybeans

To design appropriate strategies for ex situ and in situ conservation of wild soybean and safeguard the biosafety of the wild soybean gene pool when genetically modified soybeans are grown, it is important to understand its genetic characteristics, and to quantify gene flow and kinship within and between neighboring populations. We analyzed 9 pairs of neighboring populations of wild soybeans using 20 pairs of nuclear SSR markers. Results showed that Chinese wild soybean natural populations had outcrossing rates of 0–3.5% and that most populations contained many kinship families. The kinship families could be attributed to the accumulation of outcrossed offspring within populations during the history of population colonization. Wild soybean is very sensitive to environmental selection, which results in genetic differentiation of populations, and the emergence of specific alleles. We used an index τ to explain why genetic differences would exist between the pairwise populations; the interpopulation genetic differentiation chiefly consisted in the differences of allele frequencies over the genome. We found long-distance dispersal (1.5 km) of wild soybean seeds in a land ecosystem. There was close correlation between genetic and geographical distance among natural populations of Chinese wild soybean. Within a distance of 50 km, there was greater gene flow when the distance between populations was shorter. These findings have implications for ex situ and in situ conservation in an ecogeographical region, and also for protection of the gene pool from contamination by GM soybeans in wild soybean species.     

籼稻多蘖矮半矮秆基因的遗传分析和基因定位

对籼稻标记基因系材料多蘖矮的遗传分析表明, 其矮生性状是由2对隐性半矮秆基因控制的,分别为sd1和一个新的半矮秆基因,该基因初步定名为sdt3。以多蘖矮与南京6号杂交F₂的分离群体为基础,应用SSR标记进行连锁分析,将半矮秆基因sdt3定位于第11染色体的SSR标记SSR98和SSR35之间,分别相距0.06 cM、0.13 cM,二者之间的物理距离约为93kb。以南京6号为轮回亲本与多蘖矮进行回交和自交获得由半矮秆基因sdt3控制的近等基因系（新多蘖矮）,以赤霉素处理表明由sdt3控制的半矮秆系新多蘖矮对赤霉素不敏感。