物种入侵的生态学和进化生物学思考

物种入侵是指外来物种在自然状态或人类作用下,在异地生长、繁殖并建立种群的现象。目前,物种入侵已极为普遍,引起了全球性的关注。物种入侵产生了许多生态和环境问题,但是,它也为我们更好地了解自然界提供了大量的研究和实验的机会。来自于物种入侵研究(基础和应用性研究)的观点,对于理解正在形成的关于自然界的生态学、进化和生物地理学的一些观点非常重要。对于物种入侵的研究也可以支撑合理保护、管理和政策与对策的正常发展,这些对于阐述全球变化的许多方面包括物种灭绝、进化上的物种多样化以及生态系统功能都是必要的。本文从生态学和进化生物学的角度对物种入侵问题进行了分析,以期能使对物种入侵问题的认识更为全面。     

Chitosan as a biocontrol agent against the pinewood nematode (Bursaphelenchus xylophilus)

The pine wilt disease (PWD) is caused by Bursaphelenchus xylophilus and poses great environmental and economic challenges. Thus, the development of sustainable techniques for the control of this epidemic disease is of major importance. This work aimed at evaluating if the application of different molecular weight (MW) chitosans as a soil amendment could be used to control the PWD in maritime pine (Pinus pinaster, very susceptible to the disease) and stone pine (Pinus pinea, less susceptible). At the end of the experimental period (24 days after inoculation), P. pinaster and P. pinea untreated plants presented ca. 3825 ± 100 and 70 ± 47 nematodes, respectively. In P. pinaster, the high‐MW chitosan prompted the most drastic results, inducing a 21.9‐fold reduction in nematodes numbers, whereas in P. pinea, the most effective was the low MW chitosan, which reduced nematodes numbers up to 7‐fold, compared with untreated plants. P. pinea seems to be highly resistant to the disease, presenting nematode numbers up to 54.6‐fold lower than P. pinaster and less severe chlorophyll loss (ca. 2‐fold).     

Inoculation of seven pine species or hybrid seedlings with Korean isolates of pinewood nematode under greenhouse conditions

? Seedlings of seven pine species or hybrids, Pinus densiflora, Pinus thunbergii, Pinus virginiana, Pinus echinata, Pinus koraiensis, Pinus thunbergii × Pinus massoniana and Pinus × rigitaeda, were inoculated in a greenhouse with 100, 1 000 and 10 000 pathogenic nematodes, Bursaphelenchus xylophilus, to compare initial symptom development and to evaluate susceptibility to pinewood nematodes.

? Genetic types were significantly different in the starting day of needle discoloration (χ² = 43.48; P < 0.001) and the period from nematode inoculation to seedling death (χ² = 32.85; P > 0.001), but treatment groups were not statistically different in the days from inoculation to seedling death in any of the pine species.

? There was no relationship between seedling mortality and the number of nematodes recovered from infected seedlings. Only eight of the nematode-inoculated seedlings remained alive by day 83 after inoculation, including four seedlings each of P. echinata and P. × rigitaeda grown from bulked seed collections.

? Seedlings from the bulk lots were likely more genetically diverse than those from the single-source lots. A broader genetic representation of pine species or hybrids is suggested for screening for planting stocks resistant to B. xylophilus.

     

Identification and genetic analysis of the pinewood nematode Bursaphelenchus xylophilus from Pinus yunnanensis

In China, pinewood nematode (PWN), Bursaphelenchus xylophilus, was first discovered from Pinus thunbergii in 1982. Thus far, 14 species in the genus Pinus have been reported to be infected by PWN under natural conditions. Pinus yunnanensis, a pine species native to south‐western China, is considered a pioneer tree for barren hill afforestation in areas undergoing rocky desertification. In this study, we detected PWN in dead P. yunnanensis trees in Anlong County, Guizhou Province, China, using both morphological and molecular methods. To our knowledge, this is the first report of PWN from P. yunnanensis in China. To investigate the possible origin of this new outbreak, mitochondrial cytochrome oxidase gene subunit I and cellulase gene sequences were used to evaluate genetic relationships among worldwide PWN isolates. Phylogenetic tree and haplotype networks revealed that the Anlong isolate (BxChQAL008) sequence was identical to those of seven Chinese isolates collected from Sichuan, Chongqing, Zhejiang, Anhui and Shandong (372–1500 km from Anlong County), but different from the isolate BxChQZY030 collected from the same province (330 km from Anlong County). It is suggested, therefore, that more than one introduction of PWN into Guizhou Province has taken place. The Anlong isolate was likely introduced from neighbouring or more distant provinces rather than from outside China. Moreover, the absence of a correlation between geographic and genetic distance was observed using Mantel test analysis, providing evidence that human‐induced dispersal plays a fundamental role in the spread of the PWN in this region.     

松材线虫侵害马尾松林后群落谱系多样性和结构动态

调查松材线虫Bursaphelenchus xylophilus侵害马尾松Pinus massoniana林不同时间的4个群落和邻近的1个常绿阔叶林群落,利用群落净谱系亲缘关系指数(NRI)和净最近种间亲缘关系指数(NTI)等,分析松材线虫侵害马尾松林后群落谱系结构和谱系多样性动态。结果表明:松材线虫侵害后,群落物种和谱系多样性下降,侵害12 a后与常绿阔叶林的多样性最接近,侵害4,8 a群落内的物种和谱系与常绿阔叶林相似性最高,侵害0 a的与常绿阔叶林相似性最低;松材线虫侵害0,4,8 a的整个群落的谱系结构发散,侵害12 a与常绿阔叶林的谱系结构聚集。松材线虫侵害后,马尾松林群落向常绿阔叶林演替,同时,环境过滤对整个群落物种的组成起主要筛选作用。     

Pine wilt disease: detection of the pinewood nematode (Bursaphelenchus xylophilus) as a tool for a pine breeding programme

The pinewood nematode (PWN), Bursaphelenchus xylophilus, is a serious quarantine pest first detected in Portugal and Europe in 1999. It is the causal agent of pine wilt disease (PWD). A resistance breeding programme has been initiated to contribute to control the evolution of the disease. Five hundred and four adult maritime pine, Pinus pinaster, trees were phenotypically selected as candidate trees for this programme from an area affected by PWD. To identify tolerance to the nematode, the selected trees were monitored monthly. Over the course of 1 year, 57 candidate trees died and were tested for the presence/absence of the PWN. As accuracy of detection is of major importance, an ITS‐PCR‐based method applied directly to wood from adult maritime pine trees was tested and compared with a standard morphological identification method. The results showed that the use of PCR to detect the pathogen provided more rapid and accurate results in comparison with the standard morphological identification. Thus, this method is suitable to be used in the survey of the breeding population for resistance/tolerance to PWD.     

Canadian conifers as hosts of the pinewood nematode (Bursaphelenchus xylophilus): Results of seedling inoculations

Seedlings of 22 species of conifers from across Canada were inoculated with m and r form isolates of the pinewood nematode (Bursaphelenchus xylophilus). In an experiment made under ambient (summer‐fall) temperatures in a shadehouse at Victoria, British Columbia, Canada, 8 of the 22 conifer species were killed by the nematodes, but mortality was low, i.e. 4–30%. Pines (eastern white, Jack and red) were more susceptible than other conifers. Yellow cypress, eastern white cedar, western red cedar and western hemlock were not killed by the nematodes. In a second experiment made at elevated temperatures (30°C‐16h long days, 25°C‐8 h long nights) in a greenhouse, 18 of the 22 conifer species died following nematode inoculation. Again, pines (lodgepole, eastern white, western white and red) were among the most susceptible tree species and the four conifers that were unaffected in the first experiment were not killed. Tamarack and western larch, both immune at ambient temperatures, were the two most susceptible conifers at elevated temperatures. Compared to ambient temperatures, seedlings at elevated temperatures died quicker and contained more nematodes. M and r form nematodes were equally pathogenic in both experiments.     

松材线虫病与松墨天牛研究概况

松材线虫病Ｂｕｒｓａｓｐｈｅｌｅｎｃｈｕｓｘｙｌｏｐｈｉｌｕｓ是松树的毁灭性病害。最早由美国的Ｓｔｅｉｎｅｒ ,Ｑ等在路易斯安那州的沼泽松Ｐｉｎｕｓｐａｌｕｓｔｒｉｓ的木材中发现 ,后由Ｎｉｃｋｌｅ,Ｗ .Ｒ定名。但美国的研究工作开展较晚 ,是从 1979年在密苏里州发现由松材线虫造成松树萎焉后才开始研究的 (彭趋贤 1990 )。现在已有 33个州发生。世界上研究最早的是日本。矢野 190 5年首先在长崎市周围的松林中发现松树萎焉 ,之后进行了调查研究 ,1913年作了报道 ,虽然当时未能明确是松材线虫病 ,但所记述的症状实际上就是现在…     

一对姊妹线虫的杂交试验研究

基于李氏长尾线虫(Seinura liiHuang & Ye,2006)与吴氏长尾线虫(S.wuaeHuang & Ye,2006)是一对形态很相似的姊妹线虫,在原始描述之后,采用单一配对杂交方法来进一步探讨它们相互间的生殖关系。结果表明,李氏长尾线虫与吴氏长尾线虫在生殖上相互隔离,并且它们之间的生物学特性可能存在着较大差异;研究还表明,李氏长尾线虫与吴氏长尾线虫的生殖方式都为有性生殖。     

Spatiotemporal analysis of pine wilt disease: Relationship between pinewood nematode distribution and defence response in Pinus thunbergii seedlings

Pine wilt disease is of major concern as it has destroyed pine forests in East Asia and Europe. Several studies have suggested that invasion by the pinewood nematode (PWN) Bursaphelenchus xylophilus, which causes this disease, evokes an excessive defence response in pine trees, resulting in tree death. However, few studies have quantitatively evaluated the correlation between PWN distribution and tree defence responses. Therefore, the present study aimed to quantify the number of PWNs and expression levels of putative pathogenesis‐related (PR) genes in different positions of Japanese black pine (Pinus thunbergii) seedlings over time. To quantify the number of PWNs in the seedlings, we used TaqMan quantitative real‐time PCR (qPCR) assay. During the early phase of infection, most PWNs were distributed around the inoculated sites, with only a small number being detected at distant sites, but the expression levels of PR genes were highly upregulated throughout the seedlings. Both the number of PWNs and expression levels of PR genes then increased drastically throughout the seedlings, all of which exhibited external symptoms. Thus, it appears that the rapid migration of PWNs induces a defence response throughout the seedling; however, this may not be effective in controlling these parasites, thereby ultimately leading to plant death.     

Within-tree distribution and attractant sampling of propagative pinewood nematode, Bursaphelenchus xylophilus: An early diagnosis approach

Early detection is of primary importance to enable rapid actions to prevent the spread and introduction of invasive species. The pinewood nematode (PWN), Bursaphelenchus xylophilus, a serious invasive and destructive species, is listed as a quarantine pest in the legislation of more than 40 countries. However, Baermann funnel extractions of wood from discs cut from trees at breast-height often do not detect the presence of PWN in infested trees. A serious consequence of such false negatives is the loss of the best window for implementation of eradication or quarantine measures to prevent establishment of incipient PWN infestations. Here we document the within-tree horizontal and vertical distribution of PWN in infested stands in China, using a newly developed kairomonal trapping technique. Our results provide a simple, effective, rapid and non-destructive sampling method that takes into account the changes of PWN within-tree distribution in relation to pine wilt disease (PWD) symptom development. When 60–80% of the foliage has become pale green, PWN is recovered from larger diameter branches. As disease symptoms progress, PWN moves into and down the trunk. As the needles turn yellow, PWN was recovered from the trunk at 1–2 m above the ground. The correlation between the within-tree distribution of PWN and the expression of symptoms indicated a strong association between the distribution of PWN and physiological and pathological changes that develop in attacked pines through the interaction between PWN and tree. This systematic sampling technique takes into account the within-tree distribution of the nematode and should greatly enhance early detection of PWN in field surveys, monitoring and phytosanitary inspections.     

Comparison of the potential spread of pinewood nematode (Bursaphelenchus xylophilus) in Finland and Iberia simulated with a cellular automaton model

Pinewood nematode (PWN) is one of the most threatening invasive pests in the pine forests of Europe, and it has recently spread to the Iberian Peninsula via import of timber and wooden packaging material from East Asia. A cellular automaton (CA) model was developed to simulate and compare the potential spread of PWN by transportation and its vectors, Monochamus beetles in the pine forests of Finland and Iberian Peninsula. The model assumes that all pines are equally sensitive to PWN. The CA is a spatio‐temporal grid‐based model, which can easily be applied on different geographical scales. The effects of climate warming and number of entries from ports on the spread of PWN were studied. A sensitivity analysis was conducted on the most uncertain model parameters. Twenty years after hypothetical entries, the predicted area of symptomatic PWN infection (pine wilt disease, PWD) was very low in Finland compared to Iberia. This was because of the low probability of warm July in Finland. The increase in the mean July temperature increased the area of PWD‐infected pine forest relatively more in Finland than in Iberia. An increase in the number of entries also increased the area of PWD‐infected pine forest relatively more in Finland than in Iberia. The probability of PWD infection was the highest in pine forests that were close to entry points and in areas with low elevation and high human population density.     

The effect of periderm formation in the cortex of Pinus thunbergii on early invasion by the pinewood nematode

The distribution of cortical resin canals and periderm formation in the cortex of Pinus thunbergii was studied in relation to early invasion of the pinewood nematode, Bursaphelenchus xylophilus. Nematode invasion was restricted in stem cuttings of P. thunbergii in which periderm closed cortical resin canals. Early invasion of the nematodes was also restricted in stem cuttings where wound periderm had formed in response to prior nematode inoculation. It was concluded that early invasion of pinewood nematodes in living bark tissue is restricted by periderm in mature stems and by wound periderm that had formed as the result of previous nematode infection.     

Impact of the invasion of pinewood nematode and the following different removal disturbance intensities on the plant diversity of Masson pine community

Reported in this paper are our findings of the study on changes in species diversity of different Masson pine communities after the invasion of Bursaphelenchus xylophilus (Steiner and Buhren) Nickle and subsequent and different removal disturbance management approaches. Based on the results, the species diversity indices for the arbor layers of the different communities can be arranged in the following order (from high to low): 1) broad-leaved stand after the removal of all infected pine trees in the pure pine stand in Fuyang; 2) lightly infected Masson pine and Schima superba mixed stand in Fuyang; 3) uninfected stand mixture of Masson pine and Castanopsis fargesii as a control; 4) lightly infested pure Masson pine stand in Fuyang; 5) Quercus variables stand formed following the selective removal of infected pine trees from a mixed Masson pine and Q. variables stand in Zhoushan Islands; 6) pure young Masson pine stand formed following the removal of all infected pine trees from a pure Masson pine stand; 7) pure Liquidambar formosana stand after the removal of infected pine trees from a pure pine stand in Zhoushan Islands; 8) a mixed stand consisting of Pinus thunbergii and the Masson pine in Zhoushan Island; and 9) moderately infected Masson pine stand in Zhoushan Islands. All the three diversity indices (R ₀ = richness, H′ = Shannon-Wiener index, and E = evenness) for the shrub layer did not show any significant differences among the various communities except for the pure pine stand in Zhoushan Island, which had the lowest diversity. The three indices for the herb layer of the pure young Masson pine, Q. variables stand, and L. formosana stand were higher than that of other stands. The integrated analysis showed that the Masson pine forest in different geographical situations and extent of damage had distinct disparity, just as in different disturbance degrees and restoring manners. We created the “index of disturbing intensity of stump and fallen woods” (IDISFW) to represent the degree of disturbance of tree removal to plant diversity. We found that the relationship between the species diversity index and the IDISFW shows both a lower and higher degree of removal disturbance that follows the “mid-altitude bulge” theory. Specifically, both excessive and insufficient removal of infected trees will cause a decline of plant species diversity to a certain degree. Covariance analysis indicated that different IDISFW had no significant effects on the species diversity of the arbor layer, but had different impacts on that of the shrub and herb layers, and therefore can be used to assess changes in species diversity of different Masson pine communities after the invasion of pine wood nematode. __________ Translated from Chinese Journal of Applied Ecology, 2006, 17(7): 1,157–1,163 [译自: 应用生态学报]     

湿地松萎蔫病媒介昆虫松墨天牛及其携带线虫的研究

对湿地松萎蔫病媒介昆虫松墨天牛及其携带松材线虫、其它线虫的研究表明:湿地松萎蔫病木中羽化出的松墨天牛平均体长比马尾松上的短1.05 mm;从羽化开始到结束,天牛体长呈现减小的趋势,个体相差较大,体长最大的23.2 mm,最小的仅12.1 mm,相差11.1 mm,雌虫平均体长较雄虫长0.48 mm。萎蔫病木中的线虫在天牛体内表现为2种形态,线虫尾部呈指状或锥状;天牛体内的线虫有3种分布类型:所有线虫尾部为指状,所有线虫尾部为锥状,2个形态的线虫均有。湿地松松墨天牛中以尾部锥状线虫占多数,而马尾松的则以尾部指状线虫占多数。松墨天牛携带的线虫量与其体长无关,随着萎蔫病木的线虫量的增加呈上升趋势。线虫在松墨天牛体内的分布,以腹部为多,占整个虫体线虫量的75%以上,在利用松墨天牛预测湿地松萎蔫病时,取其腹部进行分离镜检即可。     

生态学中关键种与物种重要性的概念(英文)

在本文中,对关键种的概念进行了讨论,并介绍了关键种的典型特征及其在群落和生态系统中的判别。在对关键种的研究基础上,首次提出了物种重要性(SI)概念。物种重要性可以简单地理解为生态系统中物种的重要值,它由三个指数构成,即物种结构重要值(SIV)、功能重要值(FIV)和动态重要值(DIV)。本文同时评价了三大硬阔森林生态系统中乔木树种的物种重要性。