Phylogeography of Pinus tabulaeformis Carr. (Pinaceae), a dominant species of coniferous forest in northern China

How coniferous trees in northern China changed their distribution ranges in response to Quaternary climatic oscillations remains largely unknown. Here we report a study of the phylogeography of Pinus tabulaeformis, an endemic and dominant species of coniferous forest in northern China. We examined sequence variation of maternally inherited, seed‐dispersed mitochondrial DNA (mtDNA) (nad5 intron 1 and nad4/3–4) and paternally inherited, pollen‐ and seed‐dispersed chloroplast DNA (cpDNA) (rpl16 and trnS‐trnG) within and among 30 natural populations across the entire range of the species. Six mitotypes and five chlorotypes were recovered among 291 trees surveyed. Population divergence was high for mtDNA variation (G_ST = 0.738, N_ST = 0.771) indicating low levels of seed‐based gene flow and significant phylogeographical structure (N_ST > G_ST, P < 0.05). The spatial distribution of mitotypes suggests that five distinct population groups exist in the species: one in the west comprising seven populations, a second with a north–central distribution comprising 15 populations, a third with a southern and easterly distribution comprising five populations, a fourth comprising one central and one western population, and a fifth comprising a single population located in the north‐central part of the species’ range. Each group apart from the fourth group is characterized by a distinct mitotype, with other mitotypes, if present, occurring at low frequency. It is suggested, therefore, that most members of each group apart from Group 4 are derived from ancestors that occupied different isolated refugia in a previous period of range fragmentation of the species, possibly at the time of the Last Glacial Maximum. Possible locations for these refugia are suggested. A comparison of mitotype diversity between northern and southern subgroups within the north‐central group of populations (Group 2) showed much greater uniformity in the northern part of the range both within and between populations. This could indicate a northward migration of the species from a southern refugium in this region during the postglacial period, although alternative explanations cannot be ruled out. Two chlorotypes were distributed across the geographical range of the species, resulting in lower levels of among‐population chlorotype variation. The geographical pattern of variation for all five chlorotypes provided some indication of the species surviving past glaciations in more than one refugium, although differentiation was much less marked, presumably due to the greater dispersal of cpDNA via pollen.     

Paternal cytoplasmic transmission in Chinese pine (Pinus tabulaeformis)

Summary. In this paper, the stages of normal sexual reproduction between pollen tube penetration of the archegonium and early embryo formation in Pinus tabulaeformis are described, emphasizing the transmission of parental cytoplasm, especially the DNA-containing organelles – plastids and mitochondria. The pollen tube growing in the nucellus contained an irregular tube nucleus followed by a pair of sperm cells. The tube cytoplasm contained abundant organelles, including starch-containing plastids and mitochondria. The two sperm cells differed in their volume of cytoplasm. The leading sperm, with more cytoplasm, contained abundant plastids and mitochondria, while the trailing one, with a thin layer of cytoplasm, had very few organelles. The mature egg cell contained a great number of mitochondria, whereas it lacked normal plastids. At fertilization, the pollen tube penetrated into the egg cell at the micropylar end and released all of its contents, including the two sperms. One of the sperm nuclei fused with the egg nucleus, whereas the other one was retained by the receptive vacuole. Very few plastids and mitochondria of male origin were observed around the fusing sperm and egg nuclei, while the retained sperm nucleus was surrounded by a large amount of male cytoplasm. The discharged tube cytoplasm occupied a large micropylar area in the egg cell. In the free nuclear proembryo, organelles of maternal and paternal origins intermingled in the neocytoplasm around the free nuclei. Most of the mitochondria had the same features as those of the egg cell, but some appeared to be from sperm cells and tube cytoplasm. Plastids were obviously of male origin, with an appearance similar to those of the sperm or tube cells. After cellularization of the proembryo, maternal mitochondria became more abundant than the paternal ones and the plastids enlarged and began to accumulate starch. The results reveal the cytological mechanism for paternal inheritance of plastids and biparental inheritance of mitochondria in Chinese pine. Correspondence and reprints: State Key Laboratory of Plant Physiology and Biochemistry, College of Biological Science, China Agricultural University, Beijing 100094, People’s Republic of China.     

Variation in biomass and carbon storage by stand age in pine (Pinus tabulaeformis) planted ecosystem in Mt. Taiyue,Shanxi, China

Forest ecosystems play dominant roles in global carbon budget because of the large quantities stored in live biomass, detritus, and soil organic matter. Researchers in various countries have investigated regional and continental scale patterns of carbon (C) stocks in forest ecosystems; however, the relationship between stand age in different components (vegetation, forest floor detritus, and mineral soil) and C storage and sequestration remains poorly understood. In this paper, we assessed an age sequence of 18-, 20-, 25-, 38-, and 42-year-old Pinus tabulaeformis planted by analyzing the vertical distribution of different components biomass with similar site conditions on Mt. Taiyue, Shanxi, China. The results showed that biomass of P. tabulaeformis planted stands was ranged from 88.59 Mg ha^?1 for the 25-year-old stand to 231.05 Mg ha^?1 for the 42-year-old stand and the major biomass was in the stems. Biomass of the ground vegetation varied from 0.51 to 1.35 Mg C ha^?1 between the five stands. The forest floor biomass increased with increasing stand age. The mean C concentration of total tree was 49.94%, which was higher than C concentrations of ground vegetation and forest floor. Different organs of trees C concentration were between 54.14% and 47.74%. C concentrations stored in the mineral soil for each stand experienced decline with increasing soil depth, but were age-independent. Total C storage of five planted forests ranged from 122.15 to 229.85 Mg C ha^?1, of which 51.44–68.38% of C storage was in the soil and 28.46–45.21% in vegetation. The study provided not only with an estimation biomass of P. tabulaeformis planted forest in Mt. Taiyue, Shanxi, China, but also with accurately estimating forest C storage at ecosystem scale.     

华北地区油松种群遗传多样性分析

用酸性聚丙烯酰胺凝胶（APAGE）技术分析了华北地区5个油松种群在醇溶蛋白水平上的遗传多样性。95份材料分离出12条带,其中11条具有多态性,得到23种不同的带型。种群多态位点比例在16.67%～66.67%之间。Shannon信息指数和Nei’s指数的统计结果表明：关帝山油松种群的遗传多样性高于其它种群;华北地区油松种内遗传多样性为0.248 9;种群内的遗传多样性为0.150 7,占总变异的60.55%;种群间的遗传多样性为0.098 2,占总变异的 39.45%。     

中国油松林生产力格局与模拟

基于全国３８０块样地资料,从宏观上阐明了油松林生物生产力的分布格局,建立了联系叶面积指数分布规律和地植物学知识的生产力水热相关模型与地理分布模型．在油松正常分布范围内（年均气温２～１４℃,年降水量４００／３００～７００／９００ｍｍ）,林分叶面积指数随年均气温的变化呈指数递减的函数关系,这是油松林适应水分协迫环境的一种机制．     

Polysomes are associated with microtubules in fertilized eggs of Chinese pine (Pinus tabulaeformis)

Summary. Transmission electron microscopy of immunogold-labeled Chinese pine egg cells before and after fertilization revealed that polysomes are associated with microtubules (MTs) from fertilization to the 2-nucleate embryo stage. Ribosome aggregates of various size and shape were randomly distributed in the cytoplasm of the eggs before fertilization. Single MTs or clusters were observed to be free of polysomes at this stage. Upon fertilization, all polysomes were attached to MTs, and this association persisted until the formation of the polarized embryo. Thereafter, the polysomes spread into the cytoplasm and no polysome-MT association was observed in the embryo. Some of the polysomes were attached to one end of the MTs, while others appeared to form contacts along their entire length. No polysome-microfilament association was observed at any stage of the development. The polysome-MT association may provide a mechanism for MT-dependent mRNA localization in early embryo development of this plant. Correspondence and reprints: Department of Cell Biology and Physiology, University of Pittsburgh, Pittsburgh, U.S.A.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

油松胚珠发育研究中的蛋白质组技术

油松是中国特有的树种,具有重要的造林和绿化价值。但是油松胚珠败育是油松种子生产和繁殖面临的一个严重的问题,因此研究油松胚珠的发育机理和败育原因具有重要的理论和实践意义。双向电泳和质谱技术结合研究基因表达的蛋白质组技术正越来越广泛地应用到植物研究中,我们将该技术应用到油松胚珠发育的研究中,比较了两种蛋白质提取的方法,发现蔗糖提取方法比传统的TCA方法更适合于油松蛋白质的制备。将电泳分离得到的蛋白点用肽质量指纹图谱的方法进行鉴定,得到了满意的结果,优化了蛋白质质谱鉴定的条件。     

黄龙山油松林和油松+辽东栎混交林物种组成及优势种群动态

根据黄龙山油松(Pinus tabulaeformis)林和油松+辽东栎(Quercus liaotungensis)混交林共6个群丛中27个典型样地调查资料,分析了松栎林群落物种组成、多样性指数和优势种群动态.结果表明:(1)松栎混交林中灌木层和草本层植物种数均大于油松林,且灌木和草本种类因群落生境不同而有差异.(2)物种多样性分析显示,油松林和松栎混交林中由于生境不同导致灌木层物种多样性差异最大.(3)优势种群分析表明,油松林中阴坡幼苗数量较多,种群最大的为Ⅳ和Ⅴ龄级个体;松栎混交林中油松和辽东栎幼苗充足,到Ⅱ、Ⅲ龄级二者均大量减少,此后油松个体增加而辽东栎减少,说明松栎林发育过程中各层次植物物种相互制约,优势种群之间也存在竞争,且受生境条件影响较大.(4)主成分分析表明,气温、土壤、光照和总郁闭度是主要影响因子.因此,在未来森林抚育中应适当减少优势种群密度,促进异龄化;采取有效措施合理搭配灌木和草本,强化对森林发育有利的因素,提高林分质量.     

Orientation of microtubules suggests a role in mRNA transportation in fertilized eggs of Chinese pine (Pinus tabulaeformis)

Summary. Polysomes become associated with microtubules (MTs) in egg cells of Chinese pine upon fertilization, providing direct evidence for MT-based intracellular mRNA and polysome localization. We have investigated by immunoelectron microscopy the orientation and spatial distribution of MTs and their association with polysomes in the fertilized egg cells. There is a perinuclear accumulation of MTs and polysomes in the zygote soon after fertilization. At this time, some of the MTs are perpendicular to the nuclear envelope and directly connected to the outer membrane or nuclear-pore complexes (NPC) at one end, and the other ends reach to the outer tier or cortical MTs that are parallel to the long axis of the zygote. The polysomes in the perinuclear region show the same spatial and temporal pattern as the MTs. Immunolocalization of the mRNA-binding protein hnRNP indicates that the mRNAs are loaded onto the nucleus-associated MTs immediately after their export from the nuclear-pore complexes. The polysomes and mRNAs are then transported from these MTs to the outer tier and/or cortical MTs, where they further localize to the polar region of the cell. Correspondence (present address): Fengli Guo, Stowers Institute for Medical Research, 1000 E 50th Street, Kansas City, MO 64110, U.S.A.     

灵空山油松种群年龄结构与动态分析

对山西灵空山自然保护区的油松（Pinus tabulaeformis）种群进行了调查,利用生存分析理论及谱分析理论,分析了油松种群径级结构和静态生命表,绘制了存活曲线,同时分析了种群数量的周期性。主要结果如下：（1）油松种群幼苗幼树个体数较多,中径级的个体数较稳定,而高径级由于种间竞争及环境因子的影响,个体数明显下降。（2）油松种群的存活曲线呈现Deevey-Ⅱ型。动态表现为前期增长,中期稳定,后期衰退。（3）油松种群波动存在小周期,但变化不大,说明油松种群发展较为稳定。     

Regulation by light of chlorophyll synthesis in the cotyledons of Scots pine (Pinus sylvestris) seedlings

Seedlings of gymnosperms, unlike angiosperms, synthesize chlorophyll(ide) (Chl) in darkness (D). In Scots pine cotyledons ( Pinus sylvestris L.) Chl accumulation ceases in D at a low level but Chl accumulation is strongly increased by light, red light (R) being more effective than blue light (B), whereas in Pinus maritima Chi synthesis is almost light-independent. In Scots pine the capacity to form Chl can be increased by R pulses, fully reversible by far-red light, demonstrating the involvement of phytochrome. However, when B- or R–grown seedlings were transferred to D, Chl accumulation stopped immediately irrespective of the level of P_fr (far-red light absorbing form of phytochrome), indicating that the conversion of protochlorophyllide (PChl) is light-dependent. Dose response curves in R and B and simultaneous irradiation with R and B show that R and B are perceived by separate photoreceptors. The immunodetected NADPH-dependent protochlorophyllide oxidoreductase (POR, EC 1.6.99.1), assumed to regulate light-dependent Chl synthesis in angiosperms, is not correlated with the capacity of gymnosperm Chi accumulation in darkness. While two FOR bands could be separated in extracts from dark grown material (38 and 36 kDa) of Pinus sylvestris and P. maritima , only the 38 kDa band disappeared consistently in the light. However. the significance of the more light resistant 36 kDa band for chlorophyll synthesis remains unclear as well.     

施肥对油松刺槐混交林土壤微生物种群和酶活性的影响

通过对比试验研究了尿素、磷酸铵及复合肥处理对油松刺槐混交林土壤微生物种群和酶活性的影响。结果表明 :复合肥和尿素处理能够显著促进土壤微生物种群数量的增加 ,与对照相比 ,生长季平均增加了 4 2 1%和 18 4 %。复合肥有利于提高固氮菌和钾细菌数量 ,磷酸铵促进了磷细菌数量的增长。施肥处理增强了土壤酶活性 ,且季节变化明显。     

A functional-structural model for radiata pine (Pinus radiata) focusing on tree architecture and wood quality

Backgrounds and Aims

Functional–structural models are interesting tools to relate environmental and management conditions with forest growth. Their three-dimensional images can reveal important characteristics of wood used for industrial products. Like virtual laboratories, they can be used to evaluate relationships among species, sites and management, and to support silvicultural design and decision processes. Our aim was to develop a functional–structural model for radiata pine (Pinus radiata) given its economic importance in many countries.

Methods

The plant model uses the L-system language. The structure of the model is based on operational units, which obey particular rules, and execute photosynthesis, respiration and morphogenesis, according to their particular characteristics. Plant allometry is adhered to so that harmonic growth and plant development are achieved. Environmental signals for morphogenesis are used. Dynamic turnover guides the normal evolution of the tree. Monthly steps allow for detailed information of wood characteristics. The model is independent of traditional forest inventory relationships and is conceived as a mechanistic model. For model parameterization, three databases which generated new information relating to P. radiata were analysed and incorporated.

Key Results

Simulations under different and contrasting environmental and management conditions were run and statistically tested. The model was validated against forest inventory data for the same sites and times and against true crown architectural data. The performance of the model for 6-year-old trees was encouraging. Total height, diameter and lengths of growth units were adequately estimated. Branch diameters were slightly overestimated. Wood density values were not satisfactory, but the cyclical pattern and increase of growth rings were reasonably well modelled.

Conclusions

The model was able to reproduce the development and growth of the species based on mechanistic formulations. It may be valuable in assessing stand behaviour under different environmental and management conditions, assisting in decision-making with regard to management, and as a research tool to formulate hypothesis regarding forest tree growth and development.     

O_3与CO_2浓度倍增对油松针叶抗氧化酶活性的影响

以4年生的油松幼苗为试材,采用开顶箱内气体熏蒸实验,对经高浓度O_3(80nmol·mol~(-1))和CO_2(700 μmol·mol~(-1))单一及复合处理3年后,油松当年生针叶内丙二醛(MDA)含量、活性氧自由基和抗氧化酶等逆境生理指标进行了分析.结果表明:超氧化物歧化酶(SOD)、抗坏血酸过氧化物酶(APX)、过氧化氢酶(CAT)和脱氢抗坏血酸还原酶(DHAR)活性经O_3处理后显著升高,分别比对照上升了43.1%、18.9%、283.5%和142.6%,而CO_2和复合处理仅对APX和CAT活性有明显的诱导作用,但均低于O_3处理;O_3处理导致MDA含量上升51.8%,而外加CO_2则能有效逆转这一趋势;超氧阴离子(O_2~-·)产生速率和H_2O_2含量在所有处理组合间均无显著差异.上述结果说明,O_3能有效地激活油松当年生针叶内的抗氧化酶系统,而CO_2能缓解O_3所带来的不利影响,但并非通过提高抗氧化酶活性这一途径.

Abstract：

Four-year-old Pinus tabulaeformis seedlings were planted in open-topped chambers and treated with elevated O_3(80 nmol·mol~(-1)) and/or CO_2(700 μmol·mol~(-1)). After treated for three years, the malondialdehyde (MDA) content, reactive oxygen species, and antioxidant enzyme activities in the current year needles were analyzed. In treatment elevated O_3, the activi-ties of superoxide dismutase (SOD), aseorbate peroxidase (APX), catalase (CAT) and de-hydroaseorbate reductase (DHAR) increased significantly, being 43. 1%, 18.9%, 283.5% and 142. 6% higher than the control; and in treatments elevated CO_2 and its combination with el-evated O_3, only APX and CAT activities were induced remarkably but still lower than those in treatment elevated O_3. Elevated O_3 increased the MDA content by 51.8%, whereas the combina-tion of elevated O_3 and CO_2 reversed this increase efficiently. No significant differences were ob-served in the superoxide anion (O_2~-·) generating rate and H_2O_2 content among all treatments. All the results demonstrated that elevated O_3 was able to effectively activate the antioxidant enzyme system in P. Tabulaeformis current year needles, while elevated CO_2 could mitigate the negative effects of elevated O_3 through the actions other than enhancing the antioxidant enzyme activities.     

承德大窝铺不同海拔与群落类型天然油松种群遗传多样性

利用ISSR技术分析了河北承德大窝铺3个不同海拔、4个不同群落类型的油松天然种群的遗传多样性与遗传分化,探讨了天然油松种群遗传多样性与主要生态因子的关系.结果表明:天然油松种群具有较高的遗传多样性,13个筛选出的随机引物共检测出180个位点,总多态位点比率分别达到73.33％和91.11％.海拔影响油松种群的遗传多样性,中海拔( 1070～1130 m)高于高海拔(1230 ～1350 m)和低海拔(910 ～970 m)(多态位点比率分别为63.33％、51.67％和59.44％);群落类型与种群的基因组结构变异之间存在一定的内在联系.遗传变异多来自种群内部(分别为90.45％和75.22％),种群分化不明显;群落类型对天然油松种群间遗传变异和基因交流(Gst为0.248,Nm为1.517)的影响高于海拔对其影响(Gst为0.095,Nm为4.738).可以认为,群落类型对大窝铺天然油松种群遗传分化有较大影响.     

Growth space competition index(GSCI)and application in the individual intraspecies competition of Pinus tabulaeformis and Platycladus orientalis forests

dized-combination;Yield;Grey system;Related degree0玉米科学Journal of Maize Sciences50-52S513D04723D;D047了20多种衡量竞争程度的竞争指数系统,但有关学者研究后认为,这些竞争指数都没有较好地解决了定量分析中树冠变化的问题。论文首次提出生长空间竞争指数(Growthspacecompetitionindex,GSCI),系树冠表面积与胸高断面积之比,由于树冠外围面积代表树冠接受光照最强的部分并且是决定生产效率的重要指标,因此在群体和群落的竞争中作为反映竞争参数,较好地解决了定量分析中树冠变化的问题,为树冠覆盖程度的无量纲度量。本文还在油松(Pinustabulaeformis)、侧柏(Platycladusorientalis)林种内个体竞争中得到应用,与常用的简单竞争指数(Competitionindex,CI)和空间竞争指数(Growthspaceindex,GSI)进行了对比研究。     

5个油松种源光合特性的比较研究

为了揭示自然条件下不同种源油松光合特性的差异,为优良种源选择提供依据,在黄土高原种源实验地,对5个油松种源（陕西洛南LN,陕西黄陵HL,山西吕梁山LL,山西灵空山LK,辽宁千山QM）幼苗的光合生理特征及其与环境因子的关系进行了研究。结果表明：5个油松种源的净光合速率（P_n）呈双峰型,蒸腾速率（T_r）、气孔导度（G_s）亦基本呈双峰型,第1峰值均出现在上午8:00,有明显的光合午休现象。种源QM的水分利用效率（WUE）日变化呈单峰性,峰值出现在10:00,而种源LN、LL、LK和HL呈双峰型,第一峰值出现在8:00,第二峰值出现在12:00~16:00。种源LN、LL、LK和QM午间净光合速率的下降主要由气孔因素引起的,而种源HL则主要由叶肉细胞同化能力的降低引起的。种源LK的日均P_n、T_r和WUE均高于其他种源,是高光合、高蒸腾和高水分利用效率的种源,而种源HL属于低光合、低蒸腾和低水分利用效率的种源。T_a是影响各种源的P_n的首要因子,其次是VPD;RH是影响种源LN、LL和LK的T_r的首要因子;而T_a和VPD是影响种源HL和QM的T_r的主要因子。     

华北地区油松林生态系统对气候变化和CO2浓度升高的响应——基于BIOME-BGC模型和树木年轮的模拟

应用BIOME-BGC模型和树木年轮数据模拟1952-2008年华北地区典型油松林生态系统净初级生产力(NPP)动态,探究了树木径向生长和NPP对区域气候变暖的响应以及未来气候情景下油松林生态系统NPP动态变化.结果表明:1952-2008年,研究区油松林生态系统NPP波动于244.12 ～645.31 g C·m-2·a-1,平均值为418.6 g C·m-2·a-1.5-6月的平均温度和上年8月至当年7月的降水是限制该地区油松径向生长和油松林生态系统NPP的主要因子.研究期间,随着区域暖干化趋势的加强,树木径向生长和生态系统NPP均呈下降趋势.未来气候情景下,NPP对温度和降水的单独和复合变化的响应为正向.CO2浓度升高有利于油松林生态系统NPP的增加,CO2的施肥效应使NPP增加16.1％.在生态系统和区域水平,树木年轮是一种理想的指示生态系统动态变化的代用资料,可以检验和校正包括BIOME-BGC模型在内的各种生态系统过程模型.     

桥山林区油松人工林营养元素分配与积累的研究

在陕西省桥山林区3块油松人工林样地中,对油松人工林生态系统各组分中N,P,K,Ca,Mg等5种营养元素的含量,分配格局,积累规律以及它们与密度间的关系进行了研究,结果表明,油松人工林不同器官中营养元素含量排序为树叶＞树枝＞树根＞树皮＞干材;在26年生油松人工林现存量中,N,P,K,Ca,Mg总量分别为214．44,22．91,158.88,167.36,29.42kg.hm^-2,乔木层,灌木层,草本层和死地被物层养分贮量分别为498.98,3.32,7.31,83.40kg.hm^-2;油松人工林N,P,K,Ca,Mg的富集系数分别为3．98,42.8,5.69,1.08,4.01,每生产1t有机物质需要这5种元素约8.43-9.09kg.hm^-2.a^-1;乔木层中营养元素积累量为288.26-606.07kg.hm^-2,其中干村长褥铺张只累量约占乔木层的14．1％－16．8％。