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Old-Growth Platycladus Orientalis As a Resource for Reproductive Capacity and Genetic Diversity

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Old-Growth Platycladus orientalis as a Resource for Reproductive Capacity and Genetic Diversity Lin Zhu, Anru Lou* State Key Laboratory of Earth Surface Processes and Resource Ecology, College of Life Sciences of Beijing Normal University, Beijing, China Abstract Aims: Platycladus orientalis (Cupressaceae) is an old-growth tree species which distributed in the imperial parks and ancient temples in Beijing, China. We aim to (1) examine the genetic diversity and reproductive traits of old-growth and young populations of P. orientalis to ascertain whether the older populations contain a higher genetic diversity, more private alleles and a higher reproductive output compared with younger populations; (2) determine the relationships between the age of the population and the genetic diversity and reproductive traits; and (3) determine whether the imperial parks and ancient temples played an important role in maintaining the reproductive capacity and genetic diversity of Platycladus orientalis. Methods: Samples from seven young (younger than 100 yrs.) and nine old-growth (older than 300 yrs.) artificial populations were collected. For comparison, three young and two old-growth natural populations were also sampled. Nine microsatellite loci were used to analyze genetic diversity parameters. These parameters were calculated using FSTAT version 2.9.3 and GenAlex v 6.41. Important Findings: The old-growth artificial populations of P. orientalis have significantly higher genetic diversity than younger artificial populations and similar levels to those in extant natural populations. The imperial parks and ancient temples, which have protected these old-growth trees for centuries, have played an important role in maintaining the genetic diversity and reproductive capacity of this tree species. Citation: Zhu L, Lou A (2013) Old-Growth Platycladus orientalis as a Resource for Reproductive Capacity and Genetic Diversity. PLoS ONE 8(2): e56489. doi:10.1371/journal.pone.0056489 Editor: Giovanni G. Vendramin, CNR, Italy Received August 29, 2012; Accepted January 10, 2013; Published February 8, 2013 Copyright: ß 2013 Zhu, Lou. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was supported by the National Natural Science Foundation of China (project code: 31070374) and Specialized Research Fund for the Doctoral Program of Higher Education (20120003130003). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: [email protected] Introduction the genetic diversity of these trees. Beijing City has the largest number of long-lived trees in the world. In a survey of ancient and Old-growth tree populations are assumed to be potential famous trees in 2007, 28 families, 47 genera, 66 species and 39408 resources of genetic diversity and high reproductive output; old-growth trees were found, 6122 of which were older than 300 however, there is little relevant research and empirical evidence years [5], including species of Ginkgo biloba, Chinese jujube (Ziziphus for this assertion [1,2,3]. The research regarding old-growth trees jujuba), Sophora japonica, Pinus tabulaeformis, and Pinus bungeana. The has mostly concentrated on species diversity and distribution [4,5], largest number of these trees (53.8%) consists of Platycladus orientalis the relationship of the growth status and soil condition [6,7,8], (Cupressaceae), an evergreen tree species that originated in China, cultivation, maintenance and rejuvenation [9,10]. Studies of red and that can live for centuries. Natural populations of Platycladus spruce (Picea rubens) have shown positive relationships among the orientalis are rare and restricted to Beijing, Hebei, Shanxi and age, stem diameter, height and genetic and reproductive status of Shaanxi provinces [13]. However, due to its ability to thrive under old-growth and older stands of P. rubens, and old-growth forests a very wide range of climate and soil conditions, P. orientalis is an may serve as reservoirs of genetic diversity and seed sources to afforestation tree species which is widespread and naturalized maintain populations under the pressures of climate change and throughout China and as an introduced species in both North population fragmentation [1]. The gene pool of aged tree Korea and eastern Russia [13]. Previous studies on P. orientalis populations holds a variety of valuable genes, such as those for have focused on its chemical components [14] and its drought- resistance and other genes that may be very useful for sustaining resistance characteristics [15]. Direct investigation of the genetic the health of the species or artificial breeding in the face of diversity and other characteristics of old-growth and young P. environmental changes [11,12]. orientalis populations can help to determine the differences between A great number of imperial parks, tombs and ancient temples these populations and provide empirical evidence for the assertion were established in Beijing, an old city with a long history as the that the old-growth populations are genetically diverse and capital of China. These sites provide a long-term stable potential sources for obtaining seeds for conservation and environment (hundreds of years) for long-lived trees and protect reintroduction purposes. PLOS ONE | www.plosone.org 1 February 2013 | Volume 8 | Issue 2 | e56489 P. orientalis as a Resource for Genetic Diversity Here we use P. orientalis to investigate the levels of genetic polymorphic. The old-growth population in Dingling Tomb diversity and reproductive traits in old and younger artificial exhibited the highest values for the parameters of the observed populations and compare these levels to those observed in five heterozygosity (Ho = 0.805), expected heterozygosity (He = natural populations. We hypothesize that older populations will 0.880), number of alleles (Na = 15.111), number of effective harbor higher genetic diversity than younger populations. alleles (Ne = 9.586), Shannon’s information index (I = 2.291) Through this study, we will better understand the importance of and number of private alleles (PA = 9). The largest range of allele protecting old-growth trees, both for their cultural significance as a size was also detected in the 300-yr age group, the XSA population biological archive [16] and for the gene pool that they represent. (allele range = 55.889). Conversely, the lowest values for seven of Furthermore, we will understand the role of these old populations the parameters were found in the artificial 100-yr. populations in maintaining biodiversity. The objective of this study was to(1) BDC (Ho = 0.584), SGS (He = 0.635, AR = 4.213), DL (Na = compare the genetic diversity and reproductive traits (seed size and 7.222, I = 1.599, PA = 1) and DT (allele range = 30.222); the weight) of old-growth and younger populations of P. orientalis and lowest Ne was found in DJA (4.528). to (2) determine the relationships between the age of the There was no difference between the old-growth artificial tree population and the genetic diversity and reproductive traits. populations (Group 2) and natural populations (Groups 3 and 4) for the Ho, He, Na, Ne, I and AR parameters (p . 0.1). The allele Results range of Group 2 was significantly higher than that of the other three groups (p , 0.05). The Ho, HE,NA,NE, I and AR values for Reproductive Capacity Group 1 were significantly lower than those of the other groups (p Of the 241 individuals (52.28%) of the old-growth populations, , 0.05), but no difference was detected in the allele range between 126 produced cones, whereas only 69 of the 187 individuals Group 1 and natural Groups 3 and 4. The private alleles of these (36.90%) of the 100-yr. age group displayed cones. In the 300-yr. groups did not differ significantly. age group, the proportion of individuals with cones was The results of the DBH measurements indicate that values of significantly higher than the 100-yr. age group (Chi-square test; the natural old-growth populations (Group 4) were not as large as x2 = 10.047; p = 0.002). There was no significant relationship the ancient artificial individuals of P. orientalis, probably because of between the average value of the DBH of a population and the environmental stress, such as drought and competition with other percentage of individuals with cones in the artificial populations. species. No relationship was found between the DBH and genetic There were no significant differences in the seed weight between diversity parameters. In the artificial populations, conversely, the eight sampled populations (data not shown), except for the significant correlations were found between the population TMA population, which was significantly longer (0.5845 6 average values of the DBH and the average genetic diversity 0.0101 cm, mean 6 S.E.) and heavier (0.0247 6 0.0006 g, Mean parameter values of the number of effective alleles (Spearman’s rho 6 S.E.) than the other seven populations (p , 0.05). Seed length (p = 0.467; n = 16; p = 0.012), Shannon’s information index = 0.071; n = 8; Spearman’s rho = 0.667; marginally significant) (Spearman’s rho = 0.683; n = 16; p = 0.000), allele richness and weight (p = 0.037; n = 8; Spearman’s rho = 0.738) for these (Spearman’s rho = 0.567; n = 16; p = 0.002) and allele range populations were positively correlated with the average values of Spearman’s rho p the DBH (Fig. 1a, b): Older populations tended to produce heavier ( = 0.450; n = 16; = 0.015) (Fig. 2a, b, c and d). seeds. There was a strong positive relationship between the observed heterozygosity and population averages of the DBH (Spearman’s rho = 0.510; n = 16; p = 0.006) (Fig.
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