Juvenile radiata pine clonal seed orchard management in Galicia (NW Spain)

A juvenile radiata pine clonal seed orchard (Pinus radiata D. Don) in Sergude (northwestern Spain), comprising 57 clones, was evaluated for fertility, offspring diversity and genetic drift during 4 years. Flower and fruit production was used to asses clone fertility in the orchard. Fertility variation measured as “sibling coefficient” was found to be an average of 1.5, having high genetic diversity (0.99) and low coancestry (0.013). The clones varied in fertility from year to year. Effective population size in the orchard was between 35 and 40, indicating that between 60.30 and 69 %, the clones contributed effectively to seed yield. One way to improve seed yield in the orchard is by increasing the effective population size. Thinning and pruning highly reproductive clones of gibberellin application might be useful in increasing effective population size. The orchard’s parental balance showed consistent improvement over time. Cone and seed characteristics were also investigated. The results showed that significant genetic variation exists among clones for cone width and length; total, sound and empty seed number; as well as 100 seed weight. The seed orchard is still in a juvenile phase; moreover, the broad-sense heritability and variation data obtained in this study could be very valuable in the management practices of the existing radiata pine clonal seed orchard and can be also used for better planning of advanced generation of clonal seed orchards in Spain.     

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.     

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.     

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.     

Unequal deployment of clones to seed orchards by considering genetic gain, relatedness and gene diversity

Related clones in seed orchards lead to inbreeding depressionand reduced genetic value of the seeds. This study aims to developthe methodology for deployment of related clones to seed orchardswhen the breeding value and the pedigree are available for eachcandidate. The following goals were considered: high geneticgain, high genetic gain adjusted for predicted inbreeding depression(net gain), high gene diversity, and high effective clone number.The selection strategies included truncation selection withor without relatedness restrictions, maximizing genetic gain(linear deployment) with or without restrictions on relatednessand maximizing net gain. The selection strategies were appliedto Norway spruce seed orchard candidates evaluated in clonaltests. The material comprised full-sib and half-sib relationships.Comparisons were made both at the same gene diversity and atthe effective clone number. Maximizing net gain by unequal rametnumber deployment resulted in considerable higher net gain anda considerable reduction of related ramets in many comparisons.Linear deployment restricted against related clones comparedat the same status number resulted in almost as high net gain.Reduction in gene diversity may be a more important reason toavoid relatives in seed orchards than the subsequent inbreedingfor achieving a high net gain.     

Estimates of pollen contamination and selfing in a coastal Douglas-fir seed orchard

Seed orchards are the link from tree breeding to reforestation programs and are theoretically expected to function as closed, perfect populations, ensuring gain and diversity are consistently and predictively delivered as improved seed and seedlings. Seed orchard populations often deviate from panmixia due to fertility variation, reproductive asynchrony, and gene flow, leading to reduced seed crops genetic quality. Here, as a part of multiyear monitoring study, we used DNA fingerprinting (simple sequence repeat markers) to assess a Douglas-fir (Pseudotsuga menziesii) seed orchard's seed crop genetic quality (2009 seed crop). The studied seed crop was produced under ambient temperature (i.e. no reproductive phenology manipulation) and pollination was augmented by pollen from within orchard's pollen donors. DNA fingerprinting of the parental population (66 parents) along with 207 gametophyte (1n) – embryo (2n) pairs of random bulk sample of seed allowed parentage (maternal and paternal) assignment and the direct assessment of pollen contamination (0.18 ± 0.027) and selfing (0.17 ± 0.025) rates as well as parental (pollen, ovule and individual parent) gametic contribution was compared to a previous year's crop (2005). The extended reproductive phenology coupled with variable within-orchard pollen availability has created opportunities for both self and foreign pollen to be successful at various times resulting in the seemingly paradoxical scenario of high selfing and gene flow. These results (2005 and 2009) allowed comparison of seed orchard's crop management practices and are expected to provide scientific foundations to effective seed crops genetic quality improvement.     

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.     

Parental participation in progeny and effective population sizes in experimental seed orchards of wild cherry Prunus avium L. (Batsch)

Parents and progenies genetic diversity, and male and female contributions to the seed crop were assessed in three experimental Prunus avium seed orchards. Collected data were used to compare different effective population sizes, based on phenological, seed crop and paternity analysis. Our results did not show any difference of genetic diversity between parents and progenies. A limited pollen pollution was detected. We showed that distance and coflowering among clones had a significant effect on effective pollination, and a significant effect of the production of flowers was revealed in one of the seed orchards. Our study also revealed a quite low number of effective size of fathers per mother, but high effective sizes of mothers, fathers and parents at the level of the seed orchard. Finally, the calculation of effective size of mothers, fathers and parents was not highly modified when having the complete information based on the paternity analysis.     

Magnitude and efficiency of genetic diversity captured from seed stands of <Emphasis Type="Italic">Pinus nigra</Emphasis> (Arnold) subsp. <Emphasis Type="Italic">pallasiana</Emphasis> in established seed orchards and plantations

To test how efficiently plantations and seed orchards captured genetic diversity from natural Anatolian black pine (Pinus nigra Arnold subspecies pallasiana Holmboe) seed stands, seed sources were chosen from 3 different categories (seed stands (SS), seed orchards (SO) and plantations (P)) comprising 4 different breeding zones of the species in Turkey. Twenty-five trees (mother trees) were selected from each SS, SO and P seed sources and were screened with 11 Random Amplified Polymorphic DNA (RAPD) markers. Estimated genetic diversity parameters were found to be generally high in all Anatolian black pine seed sources and the majority of genetic diversity is contained within seed sources (94%). No significant difference in genetic diversity parameters (numbers of effective alleles, % of polymorphic loci and heterozygosity) among seed source categories was found, except for a slight increase in observed heterozygosities in seed orchards. For all seed source categories, observed heterozygosity values were higher (H_o = 0.49 for SS, 0.55 for SO and 0.49 for P) than expected ones (H_e = 0.40 for SS, 0.39 for SO and 0.38 for P) indicating the excess of heterozygotes. In general, genetic diversity in seed stands has been transferred successfully into seed orchards and plantations. However, the use of seeds from seed orchards can increase the amount of genetic diversity in plantations further. The study also demonstrated that number of plus-tree clones (25–38) used in the establishment of seed orchards was adequate to capture the high level of diversity from natural stands.     

Variation in effective number of clones in seed orchards

The effective number of clones (N _c) wasestimated for 255 conifer clonal seed orchards in Finland, Korea, andSweden, based on the variation in the number of ramets among clones. Themean census number of clones (N) varied from 70, in 13 KoreanPinus koraiensis seed orchards, to 139 in 176 Finnish Pinussylvestris seed orchards. The mean effective number of clones(N _c) was 66, with a range from 10 to 421. One fifthof the orchards had N _c between 10 and 40, and twothirds between 41 and 160. On average, the relative effective number ofclones (N _r =N _c/N) was 0.74, with a range from 0.2to almost 1.0. Thus, the census number of clones in a seed orchard isgenerally rather informative, but the effective number of clones is moreinformative. Many of these first-generation seed orchards wereestablished with an intention to have near-equal numbers of ramets foreach clone. The use of effective number of clones may be more importantin future seed orchards and genetically thinned seed orchards.     

Background pollination in Pinus sylvestris seed orchards

The proportion of background pollen grains in the seed crops of two Pinus sylvestris seed orchards in Central Finland was estimated with the aid of multilocus allozyme markers. The orchards studied were more than 30 years old and in full pollen production. For the bulked seed crops of the seed orchard with southern clones the estimated average of background pollination over four years was 26%. There were statistically significant differences between years. No significant heterogeneity in the degree of background pollination between clones was found. Among single ramets there was significant heterogeneity in the estimated contamination rates, but the variation was not related to position in the orchard. For the seed orchard with northern clones the bulked seed crop was studied only for one year and the level of background pollination was found to be 33 %. These estimates are fairly high, but lower than for many other orchards. Background pollination at this level will cause losses in expected genetic gains. Part of the seeds from northern orchards will not be adapted to the intended area of use.     

Genetic thinning of clonal seed orchards using linear deployment may improve both gain and diversity

The linear deployment algorithm defines a straight line relationship between the number of ramets and the breeding value of a given clone. However, when used to determine thinning in clonal seed orchards, there is an upper limit, determined by the number of ramets in the seed orchard prior to thinning. Linear deployment thinning maximises genetic gain and effective clone number whilst minimising thinning intensity, thus both production and genetic diversity are optimised. Here two applications of linear deployment to Swedish Norway spruce seed orchards are described.     

Variation in cone and seed characters in clonal seed orchards of <Emphasis Type="Italic">Pinus sylvestris</Emphasis>

Cone and seed characters were observed on top, middle and bottom portions of tree crown in 3 ramets/clone in three Turkish Pinus sylvestris seed orchards. Broad sense heritability (clonal repeatability), and correlations among characters were estimated. Around one quarter of the seed production occurred in the top portion, half in the middle and one quarter in the bottom portion of the crown for all orchards. The percentage of filled seeds varied little with the crown position, indicating more or less similar levels of selfing in the bottom of the crown as in the top. The seed weight was typically 11 mg. Differences were found for studied cone and seed characters among orchards and crown positions. Variation among grafts within clone was higher than among clones for most characters. The heritability was on average below 0.5 (e.g., cone diameter, number of filled seed per cone) and rarely rose above that (e.g., cone form, length/diameter; percentage of filled seed) for any individual characters. The coefficient of variation within clones was often higher than among clones. Thus, non-genetic factors seem often more important for the variation in performance of grafts than their genetic constitution. Cone form (length/diameter) was the character where the clone influence was the strongest. Cone number and cone dry weight showed significant correlations with seed characters (numbers of total and filled seed, percentage and weight of filled seeds). Significant correlation was found between seed characters.     

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.     

马尾松种子园优良无性系的选择与评价

通过对马尾松种子园132个无性系种子产量性状调查与木材比重测定及种子园自由授粉子代8年生生长性状调查,在统计分析、相关分析基础上采用综合评分法进行优良无性系的选择,入选率为20%。结果表明:对马尾松种子园无性系再选择,以营建改良代种子园是可行的。8年生子代材积可获得改良增益10.6%,木材比重可获得4.6%的增益,种子园单株产种量可获得增益37.5%,选择效果明显,达到提高种子园产量、增加改良增益的目的。     

促进油松种子园结实技术研究

去劣硫伐是提高种子园产量的主要措施之一，而无性质再选择能科学地确定疏伐对象。通过子代测定和对无性系结实量观测，采用综合指数法综合评定出优、中、劣无性系，从而确定疏伐对象、疏代强度、疏伐时间等。对已进入盛果期的种子园可对结实母树进行整形修剪，增加结实量。施肥能促进种子园母树生长和开发结实，提高种子产量。     

樟子松无性系种子园种子生产及其苗木培育

樟子松种子园无性系间的花粉量、种子产量均存在着差异,这种差异,是由樟子松无性系本身的遗传基因决定的。种子园中,往往少数无性系控制着园内50%以上的花粉量和种子产量。由于部分无性系种子产量低,育苗、造林时这些家系容易丢失。为减少家系的丢失数量,育苗时,应改善圃地的环境条件,增加土壤肥力,加强除草等抚育措施,提高育苗水平。     

Allozyme frequencies,outcrossing rate and pollen contamination in Picea abies seed orchards

Allozymes of Swedish seed orchard clones of Norway spruce (Picea abies Karst.) were studied. There were differences between gene frequencies of origins from different regions. The gene pool of the fertilizating pollen of two different orchards was different. Pollen contamination and outcrossing rates were estimated in two seed orchards. The fraction of paternal gametes which did not originate from any seed orchard clone was 10% and 17%, respectively. Based on simulation it was estimated that only a quarter of the gametes with origin outside the orchard would be detected. Estimation of multilocus outcrossing rates were 0.95 and 0.98.     

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.     

Variability in seed traits and genetic divergence in a clonal seed orchard of Dalbergia sissoo Roxb.

The variations in seed and pod traits, genetic superiority and genetic divergence were evaluated for a Clonal Seed Orchard (CSO) of Dalbergia sissoo Roxb. at Bithmera, India consisting of 20 clones from different agro-climatic conditions of four northern states (Uttar Pradesh, Uttarakhand, Haryana and Rajasthan). The seeds and pods of various clones in the orchard exhibited significant variability in size, weight and other characters. Significant positive correlations were observed between seed length and seed width (p<0.05), seed length and seed thickness (p<0.01), seed length and seed weight (p<0.01), seed thickness and seed weight (p<0.01), seed length and germination value (p<0.05). The genetic parameters for seed and pod traits also showed a wide range of variations in the orchard. Heritability values were found to be over 50 percent for most of the seed and pod traits. Seed weight, seed length and seed thickness showed high heritability values coupled with maximum genetic gain for these characters. Ward’s minimum variance dendrogram of clones of D. sissoo showed three distinct clusters; cluster 1 was the largest with 12 better clones whereas cluster 2 and 3 consisting of seven moderate clones and one poor clone, respectively. Mean cluster values showed sufficient variation among the clusters for seed weight, germination value and seed length. The possible hybridization between best clones of cluster 1 to the disease resistant clone of cluster 2 (resistant against deadly Gandoderma lucidum root rot disease of D. sissoo) is also suggested for further breeding programmes of the species. The deployment of clone 194 (better performed and disease resistant) is also recommended in future plantation programmes of D. sissoo in northern India.