Female fertility variation in mature Pinus sylvestris clonal seed orchards

Abstract

Female fertility is the basis for the output of seeds from clonal orchards and its variation is of major interest for the economics and biology of seed orchards, especially for the efficiency and diversity of seed orchard crops. Assessments of female fertility variation in 10 mature (>15 years old) seed orchards of Scots pine (Pinus sylvestris L.) were evaluated and compared. Depending on the individual orchard, fertility variation for each clone was assessed in slightly different ways, e.g. number of strobili, cones, seeds or litre of cones per ramet. Assessments in five of the orchards were made over consecutive years. The main result was that the clonal variation in mean female fertility per surviving ramet was lower than expected from the literature; the Kang–Lindgren sibling coefficient (Ψ) within individual years averaged 1.35. The variation between ramets within clones and years was of similar magnitude as that between clones. Clone by year interactions were noticeable, but were slightly lower than the variation between as well as that within clones in individual years. There was considerable variation in the variance components between years. The limited variation in female fertility indicated that it should not be a selection criterion when selecting clones for a seed orchard. Furthermore, it will not result in large differences between clones in seed set or large reductions in gene diversity in productive Scots pine seed orchards.     

Impact of Fertility Variation on Gene Diversity and Drift in two Clonal Seed Orchards of Teak (Tectona grandis Linn. f.)

Two 25 year old teak clonal seed orchards comprising 15 (CSO-I) and 20 clones (CSO-II), respectively, selected mostly from moist forests of Western Ghats (latitude 10° N) in southern India, were evaluated for fertility, offspring diversity, and genetic drift. The orchards differed in fertility of clones as well as flower and fruit production per ramet. Fertility was highly skewed in CSO-II, where one clone (originating from higher latitude −17° N, in Eastern Ghats of peninsular India) produced 55% of the fruits and 68% of the flowers in the orchard, in contrast to a similar contribution from four most fertile clones in CSO-I. Fertility variation, measured as `sibling coefficient' (1.7 in CSO-I and 8.3 in CSO-II), was high in CSO-II resulting in high coancestry and low effective population size (3 times lower than CSO-I) in the seed crop. In CSO-I, 58% of the clones contributed effectively to seed production compared to only 12% effective contribution resulting in eight times higher genetic drift in CSO-II. Placing limits on how much seed can be collected per clone might be useful in restricting over representation of highly reproductive clones thereby increasing genetic diversity in the seed crop.     

Clonal variations in nutritional components of <Emphasis Type="Italic">Pinus koreansis</Emphasis> seeds collected from seed orchards in Northeastern China

From four Korean pine (Pinus koraiensis) orchards, 60 clones were selected and analyzed for the fatty acid and amino acid components of the seeds to reveal the variations and correlations of the seed characteristics among the clonal source orchards and clones. The nutritional components of the seeds of the P. koraiensis trees exhibited rich genetic variation; the variation coefficient of the fatty acids was 2.24–66.83 %, while the variation coefficient of the amino acids was 14.70–38.88 %. Relatively high genetic-improvement potential exists for the nutritional components of the seeds. The phenotypic differentiation of the fatty acid and amino acid components reveals that variation within the population (85.18 %) was the primary source for the variation of the fatty acid components; variation among the orchards (63.08 %) was the primary source of the variation of the amino acid components. Data drawn from various clonal source orchards all showed that the seed characteristics were highly controlled by heritability (h² > 80 %), and the seed characteristics of the P. koraiensis trees exhibited a similar genetic gain trend. The principal components were analyzed to obtain the comprehensive principal component value for each clonal seed orchard. Twelve clones were selected based on a clonal selection rate of 20 %. Correlation and multiple stepwise-regression analyses were conducted, considering different location conditions, to reveal the stable correlations between the seed characteristics to facilitate improvements of the seed yield of P. koraiensis trees and the clonal selection. Species of real characteristics in P. koraiensis were controlled by higher heritability. Genetic gain was obtained by selecting of superior clones.     

Clonal variation in chemical wood characteristics in Hanönü (Kastamonu) Günlüburun black pine (Pinus nigra Arnold. subsp. Pallasiana (Lamb.) Holmboe) seed orchard

ABSTRACT

Scientific evidence suggests forest cover across the world is gradually decreasing while the world’s population is rapidly increasing. In line with this, the demand on raw wood is also increasing, making it impossible for the already limited natural resources to meet human needs in proportion to the increasing population size. To counter this, there is a need to increase the variety and availability of timber through genetic improvement. This has been seen in genetic improvement studies in forestry which have resulted in seed orchard establishments; however, the genetic variation in these seed orchards continues to be a topic of debate. This study aims to determine the clonal variation in chemical characteristics in a Hanönü (Kastamonu) Günlüburun black pine (Pinus nigra Arnold. subsp. pallasiana (Lamb.) Holmboe) seed orchard. The analysis shows differences between the clones were seen at the 99.9% confidence level. Ash content, NaOH, extractive substance, and hot water solubility exhibited the highest heritability scores, according to the heritability estimates of the clones in the seed orchard. However, hemicellulose content showed an extremely low H₂ (0.1).     

马尾松种子园无性系结实量、种实性状及遗传参数的分析

马尾松(Pinusmassoniana)是我国南方重要的用材树种。80年代初在马尾松主产区建立了一批初级无性系种子园,并开展了种子园有关营建技术的研究。反映种子园无性系繁育能力的种实性状包括了球果及种子性状,是林木种子园经营管理者所关注的重要内容,它涉及到种子园的产量和种子品质等问题。国外对火炬树(P .teada)、阿勒皮红松(P .abasica)、斑克松(P .banksiana)、白云杉(Piceaglaua)、北美黄杉(Pseudosugamenziesii)等树种(Matthewsetal.,1986 ;Matzirisetal.,1998;Todhunteretal.,1981;Horh ,1984 ;Clairetal .,1991) ,研究了花粉产量…     

Genetic variation in fruitfulness in a hinoki (Chamaecyparis obtusa Endl.) seed orchard and its impact on the maintenance of genetic diversity in seedlots

Genetic variation in seed/cone production among clones was studied in a hinoki (Chamaecyparis obtusa Endl.) seed orchard containing 25 plus-trees by analyzing the number of cones, the yield of cones and seeds of individual ramets for 5 successive years (1982 to 1986). There was significant variation among clones each year and parental contribution in the seed orchard. Specifically, in the years 1982, 1983, 1984, 1985, and 1986, 20% of the clones produced 37.2, 60.6, 36.0, 44.3, and 44.8% of the total cones, respectively. The size of the crop greatly influenced the parental balance in the resulting seed/cone crops. The product moment correlation coefficients and Spearman’s coefficients of rank correlation were small and insignificant between consecutive years, but large and highly significant between alternate years, suggesting the presence of carry-over effects in seed/cone production. The broad-sense heritability on a clone mean basis was 0.74 ± 0.15 for the number of cones, 0.72 ± 0.14 for the yield of cones, and 0.68 ± 0.13 for the yield of seeds. The corresponding heritabilities from analyses combined over all years were 0.24, 0.558, and 0.724, respectively. These results indicate that seed/cone production in hinoki is under strong genetic control. Several managerial measures are discussed that maintain the genetic diversity in seedlots used for reforestation, by reducing the variation in seed/cone production among clones and producing seed crops with equal contributions from all parents.     

Genetic variation, mating patterns and gene flow in a Pinus pinaster Aiton clonal seed orchard

? Relatedness among parents, variation in clonal fertility and background pollination deviate the realized genetic gain and the gene diversity of open pollinated seed orchard from expectation, in particular in wind pollinated species such as Pinus pinaster Aiton.

? This work investigates the genetic variation, the mating system and the pollen contamination in a P. pinaster clonal seed orchard (CSO), by screening the 60 clones from the CSO and the seeds collected from 21 mother-trees with three nuclear microsatellites.

? The expected diversity was similar, but the observed heterozygosity decreased 20% in the progenies compared with the parental trees. The outcrossing rate was 90.1%, the biparental inbreeding 21.7% computed through a multilocus approach, and the observed selfing 3.9%. The observed gene flow from outside the CSO was 52.4%.

? From the results we concluded that the observed gene flow and the biparental inbreeding were high, and care should be taken in the implementation and management of future CSO, in particular clones should be checked for relatedness and the ramet number could be directly proportional to their breeding value.

     

气候条件对马尾松种子园产量的影响

对信宜马尾松种子园的85个无性系进行种子产量与花期和气候因子的关系分析。结果表明：雌花始花期较早的无性系种子产量较高；气候条件对种子产量影响很大，授粉期间旬平均温度及旬积温较高，湿度较低，没有明显的倒春寒时对种子产量有利。在保证具有优良遗传品质的前提下，可选择雌花始花期较早的无性系作为建园亲本。     

Temporal and spatial variation of flowering among Pinus nigra Arn. clones under changing climatic conditions

Until recently, the most important factors affecting the economics and genetics of the seed crop from seed orchards were considered to be the timing and duration of flowering, variation in fertility and the total number of clones used to establish the seed orchards. Change in climatic conditions however is an emerging factor that could prove crucial regarding the timing of flowering and synchronization among clones and thus, the quality and quantity of seed production. The temporal and spatial variation in flowering phenology and the duration of flowering were studied in consecutive years in a Pinus nigra Arn. seed orchard. Sixty plus trees representing the distribution of the species in Northern Greece were used to establish the seed orchard, and nineteen ramets per clone were planted in a honeycomb experimental design in order to avoid kinship. Temporal variation among clones, as well as spatial variation among ramets within clones growing at different sites of the orchard were recorded, for initiation and duration of male and female flowering. The majority of clones were synchronized in dates of flowering during the year with weather conditions close to the long-term climatic conditions, except for a limited number of clones that were precocious or late flowering. The pronounced variability in climatic conditions over the 2 years strongly affected the flowering and synchronization among clones, resulting in almost complete asynchrony during the xerothermic year, which was characterized by a prolonged mean monthly temperature increase of 2.3 °C and a water deficit of 53% in a 7-month-period (November to May). These results suggest that one of the effects of a warmer and drier climate may be the lack of flowering synchronization, as pollen shedding might be completed before female conelets reach the phase of receptivity. The restriction of male parentage to a limited number of clones severely violates the panmixia assumption and could result in fertilization failure. The projected climate change for the Mediterranean region could potentially prove detrimental for fertility and flowering synchronization of forest trees, having consequences on the quantity and genetic diversity of the seed crop in seed orchards, and the natural regeneration of forest trees in forest ecosystems due to the reduced percentage of sound seed.     

马尾松种子园及其附近人工林的亲子代群体遗传结构分析

利用同工酶标记对广西覃塘林场马尾松无性系种子园及其附近一人工林的亲子代群体遗传结构进行研究。研究结果表明，不论是种子园还是附近人工林均表现为：亲代群体处于遗传平衡状态，杂合体相对过量，但子代已偏离遗传平衡，杂合体相对不足，亲子代的基因频率变化不大。种子园亲代群体的遗传多样性略高于子代，而附近人工林的亲代群体遗传多样性略低于子代。本文研究的种子园具有较宽的遗传基础，该种子园附近人工林的遗传多样性与种子园接近，但不论是人工林本身还是它的子代其多样性平均水平都较种子园的亲代和子代略低。     

马尾松二代无性系种子园的花期同步性

【目的】研究马尾松二代种子园无性系的开花物候特征、花期同步性、结实状况及气候影响因子,评价无性系花期同步性规律,指导种子园经营生产及提高种子遗传品质与产量。【方法】连续2年对浙江省淳安县姥山林场马尾松二代无性系矮化种子园的22个无性系进行花期、花量和结实性状调查,分析其开花物候与花期同步指数。【结果】各无性系雌雄球花整体花期持续时间为8～17天, 2014年始花期相对于2013年滞后约6天,且持续时间缩短约6天。整体水平上,年度内雌球花及雄球花的各花期时长差异较小;无性系的始花期和盛花期的物候特征年度间存在差异, 2014年无性系内雌雄球花开花的同步性更强。无性系的开花物候受气候因素影响较大,这使得无性系间开花同步程度降低,花期同步指数变异增加, 2013年和2014年花期同步指数变化范围分别为0.041～0.556和0.284～0.802,无性系的平均变异系数分别为32.3%和16.8%。各无性系分别作为父、母本时,年度内无性系花期整体同步性水平差异较小,表现出无性系作为父本时的散粉期整体的同步性要高于作为母本时的可授粉期的同步性。年度间无性系作为父、母本及自交的花期同步指数均有极显著差异, 2014年各无性系的花期同步指数在整体水平均高于2013年,但自交指数在年度间无显著差异。相关分析表明,雌球花越多的无性系的球果数越多;而雌球花花期持续时间越长的无性系座果率越高;不同年度的雌球花盛花期和花期的持续时间均与千粒质量指标呈显著或极显著正相关。【结论】浙江姥山林场马尾松二代无性系种子园雌、雄球花的花期具有较好的同步性,无性系间没有出现花期不遇现象。无性系交配组合的雌球花可授粉期和雄球花的散粉期影响了种子园无性系的种实品质及座果率。连续2年的观测表明无性系间花期整体同步性水平存在差异,且这种差异与二代种子园无性系的亲本遗传背景有关。各无性系分别作为父、母本时,表现出无性系作为父本时的散粉期整体的同步性要高于作为母本时的可授粉期的同步性,无性系作为父本和母本时花期同步指数呈负相关,且种子园内存在1/3无性系具有高于异交平均水平的自交可能性,是马尾松二代种子园管理和无性系再选择应综合考虑的因素。     

Mating patterns,genetic composition and diversity levels in two seed orchards with few clones—Impact on planting crop

Paternity analyses using microsatellites were conducted in two conifer clonal seed orchards (CSOs). The aim was to study mating patterns and potential dysfunctions such as unbalanced male contribution, and how these factors influence genetic diversity. The effect of unequal male contribution was quantified by the status number (N_S), which reflects build up of coancestry in the seed orchard crop due to low number of clones and unequal male contribution. This approach was also used to quantify the effect of clonal differences in germination percentages on effective population sizes. Furthermore the genetic diversity of the Abies alba and Larix kaempferi CSOs was compared to other alternative seed sources via observed heterozygosity and allelic richness.     

努力为繁荣我国林木育种事业作贡献——考察广东三个林木良种基地的点滴感受

文章扼要介绍了英德、龙山和信宜三个良种基地建设情况,提出良种生产单位的首要任务是不断给林业生产单位提供品质愈来愈好的种苗,这同时也是自身生存发展的条件;加大育种资源的研究利用,可以采取“先易后难”、“优者居先”,逐步开展,力争取得实效;重视种内各层次的遗传变异,要在地理变异的基础上,不失时机地抓紧子代测定,为了取得实效,应重视亲本组合的选择;提高种子园种子产量,增加无性繁殖系数是当务之急;加强领导,完善管理、生产与研究三结合是良种基地持续发展的条件。     

马尾松第二代无性系种子园营建技术试验研究

介绍了福建省顺昌埔上国有林场和邵武故县国有林场马尾松第二代无性系种子园建立技术和管理技术措施。通过4 a的实施,埔上国有林场和故县国有林场分别建成了面积为7 hm2和8 hm2的马尾松第二代无性系种子园。埔上国有林场建园的103个无性系嫁接后2.5 a平均抽高达184.8 cm,平均嫁接保存率达73.03%,长势良好;故县国有林场建园的97个无性系嫁接后1 a平均抽高达60.9 cm,平均嫁接保存率达94.0%,长势良好。文中还对今后的经营管理提出了方向。     

Genetic diversity of Pinus densiflora pollen flowing over fragmented populations during a mating season

We attempted to evaluate the genetic diversity of long-distance transported pollen flowing over fragmented Pinus densiflora populations during a mating season. A P.?densiflora clonal seed orchard, which was located in a rural area where many fragmented populations exist, was selected for pollen capture. Immigrant pollen captured by three clones having different flowering times was regarded as the pollen flowing over fragmented populations during a mating season. The genetic diversity (H _e) values of the immigrant pollen captured by the three clones were high (H _e?>?0.894). The correlation of paternity (r _p) values of the seeds having immigrant parent generated from the three clones were calculated to be negative. From these parameters, the pollen cloud is considered to have maintained high genetic diversity during the mating season. The genetic composition of the pollen cloud showed slight variation. The pollen captured by different trees (i.e., clonal ramets of the three clones) was significantly different based on analysis of molecular variance. Especially, the pollen pools captured by trees planted in the western side of the orchard were significantly different from the gene pool of the surrounding populations. Factors affecting this differentiation could be that the donors of the pollen transported to the orchard vary with time, as well as nonuniform dispersal of the pollen. From these results, the pollen flowing over fragmented P.?densiflora populations is considered to have high genetic diversity, compensating to some extent for fragmentation.     

Balancing seed yield and breeding value in clonal seed orchards

Seed orchards should produce seeds that are both abundant and of high genetic value. This study suggests methods to achieve such a compromise and study their efficiency. The methods were applied on data obtained from 41 seed orchard clones of Scots pine from mid-Sweden. The value of the seed orchard crop was set as a function of its breeding value, the amount of seeds produced and their gene diversity, measured as the effective number of clones. The proportion of ramets of different clones that maximized this value was regarded as the optimum for deployment of the clones in a seed orchard. The results were compared with truncation selection for breeding value, truncation selection for clone benefit (the product of seed production and breeding value) and linear deployment (where ramets are deployed linearly in relation to breeding value). The influence of two parameters was studied: the relative importance of breeding value for seed value and the size of the penalty for reducing the value of the seed crop with respect to lost gene diversity. The conventional wisdom is to select the clones with the highest breeding values, but that turned out to be the most inferior alternative studied. Clone benefit truncation provided a good approximation to optimal benefit for cases, where the effective number was low and dependence of breeding value limited.     

湿地松改良种子园部分无性系的早期遗传评价

利用湿地松改良种子园的建园无性系生产的8个全同胞家系和20个自由授粉家系苗木共营建测定林3块。3－4年生的测定林数据分析结果表明，参试的自由授粉家系作全同胞家系的平均生长量显著地大于台山湿地松初级种子园的，20个自由授粉家系的树高、胸径、材积平均增益分别为5．54％，7．77％和23．41％，初步实现了建园目标；同一无性系的不同采种年份的自由授粉子代生长存在着差异，有随着种子园母树年龄的增大而表现渐佳的趋势；改良园中还存在着少量的误选无性系，需要继续作测定和淘汰。从无性系亲子的生长量相关分析中发现，生长较差的自由授粉家系，较大部分产自生长量缩小的母本无性系。     

湿地松种子园无性系间开花结实差异分析

湿地松种子园内无性系间的生长，开花和结实等性状存在着显著差异，无性素水平和单株水平上的性状相关分析结果基本一致，雌花量与结实量，球果长与宽有较密切的相关关系，而无性素的雌花量与雌花量存在负相关关系。     

马尾松种子园营建初报

于宜昌市林科所营建马尾松种子园20hm~2。其中无性系园(马尾松一代重建嫁接种子园)13.333 hm~2,共收集154个经过子代鉴定、遗传增益大于10%的优良无性系;实生苗种子园6.667 hm~2,于枝城市、远安县、咸丰县等马尾松优良种源区选择优树145株,选用了120个优树家系苗。建园中采用了圃地小砧嫩枝诱根嫁接、一次性定植、计算机软件(SODP)优化排列设计等先进技术。     

Genetic divergence for growth and wood parameters in different clones of Dalbergia sissoo Roxb

The wood analysis for different parameters was carried out in a clonal seed orchard of Dalbergia sissoo Roxb. established during 1997 at Hoshiarpur, Punjab, India. Twelve clones with higher index value were subjected to Euclidean Cluster Analysis based on wood and growth parameters to group into seven clusters. Cluster I and II contained four and three clones, respectively, and remaining clusters had just one clone each. Clone originated from Barielly, Uttar Pradesh of cluster VII was found to be the most divergent clone. Cluster II with three clones maintained greater inter-cluster distance with other clusters. The divergence analysis has confirmed that the clones planted in the clonal seed orchard are sufficiently divergent and seed harvested from the orchard would maintain high diversity.