日本落叶松人工林种群自然稀疏规律的研究

一、前言日本落叶松(Larix kaempferi)原产日本。种群地理分布范围为北纬35°20′—38°10′,东经136°45′—140°30′,海拔高度为1000—2800m。1900年引入我国,以其较强的适应性“定居”下来,而且生长速度在抚顺地区远比长白落叶松(L.olgensis var.changpaiensis)、兴安落叶松(L.gmelinii)为快;我国黑龙江、吉林、辽宁、山东、山西、陕西、北京、河南、河北、江西、湖北、四川以及新疆等地都有人工栽培,而以吉林、辽宁东部栽培面积较大,在年平均气温2.5—12.0℃、年降水量500—1400mm 的水热条件下能正常生长。     

剪叶损伤与昆虫取食对兴安落叶松（Larix gmefinii）针叶中缩合单宁诱导作用的差异

单宁是植物内与抗虫性相关的重要次生代谢物质。为了研究昆虫取食及剪叶诱导对兴安落叶松（Larix gmelinii）针叶内缩合单宁含量的影响,用剪叶法和落叶松毛虫（Dendrolimus superans）幼虫取食处理5年生兴安落叶松苗,以香草醛．盐酸法测定其剩余健康针叶内缩合单宁含量的变化。结果表明：（1）处理后1～10d,剪叶4枝75％,8枝50％、75％,12枝25％、50％、75％及虫伤4枝75％的兴安落叶松健康针叶内的缩合单宁含量均明显高于对照（P〈0．01）,15d时,剪叶8枝75％,12枝50％的兴安落叶松针叶内的缩合单宁含量仍高于对照（P〈0．05）,说明剪叶与虫伤处理均能诱导缩合单宁含量增加。（2）剪叶4枝75％,8枝50％,12枝25％、75％的处理之间,诱导的缩合单宁含量差异不显著（P〉0．05）。剪叶8枝75％,12枝50％诱导缩合单宁含量增加幅度比其它剪叶处理大,且诱导持续时间长。说明剪叶程度虽能影响缩合单宁含量,但二者并不呈线性关系。（3）剪叶4枝75％在5d时诱导缩合单宁含量最高,虫伤4枝75％在10d时诱导缩合单宁含量最高,说明剪叶诱导的缩合单宁含量高峰早于虫伤处理。但处理后1-10d,剪叶及虫伤4枝75％的落叶松针叶内缩合单宁含量均明显高于对照（P〈0．01或P〈0．05）,之后与对照差异不显著（P〉0．05）,说明在受到诱导处理后,兴安落叶松产生应激反应,使其针叶内缩合单宁含量在一定时间内先增加,后逐渐恢复到正常水平。由上可见,可以采取适当的损伤处理取得与昆虫取食相似的兴安落叶松抗性反应。     

剪叶损伤与昆虫取食对兴安落叶松(Larix gmelinii)针叶中缩合单宁诱导作用的差异

单宁是植物内与抗虫性相关的重要次生代谢物质.为了研究昆虫取食及剪叶诱导对兴安落叶松(Larix gmelinii)针叶内缩合单宁含量的影响,用剪叶法和落叶松毛虫(Dendrolimus superans)幼虫取食处理5年生兴安落叶松苗,以香草醛-盐酸法测定其剩余健康针叶内缩合单宁含量的变化.结果表明:(1)处理后1～10d,剪叶4枝75%,8枝50%、75%,12枝25%、50%、75%及虫伤4枝75%的兴安落叶松健康针叶内的缩合单宁含量均明显高于对照(P<0.01),15d时,剪叶8枝75%,12枝50%的兴安落叶松针叶内的缩合单宁含量仍高于对照(P<0.05),说明剪叶与虫伤处理均能诱导缩合单宁含量增加.(2)剪叶4枝75%,8枝50%,12枝25%、75%的处理之间,诱导的缩合单宁含量差异不显著(P>0.05).剪叶8枝75%,12枝50%诱导缩合单宁含量增加幅度比其它剪叶处理大,且诱导持续时间长.说明剪叶程度虽能影响缩合单宁含量,但二者并不呈线性关系.(3)剪叶4枝75%在5d时诱导缩合单宁含量最高,虫伤4枝75%在10d时诱导缩合单宁含量最高,说明剪叶诱导的缩合单宁含量高峰早于虫伤处理.但处理后1～10d,剪叶及虫伤4枝75%的落叶松针叶内缩合单宁含量均明显高于对照(P<0.01或P<0.05),之后与对照差异不显著(P>0.05),说明在受到诱导处理后,兴安落叶松产生应激反应,使其针叶内缩合单宁含量在一定时间内先增加,后逐渐恢复到正常水平.由上可见,可以采取适当的损伤处理取得与昆虫取食相似的兴安落叶松抗性反应.     

中国东北落叶松属植物rbcL 基因的序列分析及系统演化

长白落叶松的分类位置及与兴安落叶松的种间关系, 长期以来未有定论。本研究从大兴安岭、小兴安岭及张广才岭等6 个地点采集实验材料, 对其叶绿体DNA 的rbcL 基因进行PCR-RFLP 分析并测序。结果显示它们的rbcL 基因序列均没有差异, 表明东北地区落叶松属植物有共同起源。在系统演化上, 长白落叶松来自兴安落叶松(Larix gmelinii Rupr.), 应作为兴安落叶松的变种即L.gmelinii Rupr.var.olgensis Ostenfeld et S.Larsen     

长白落叶松氮素营养及与生长的关系

不同季节测定了不同种源长白落叶松针叶中硝酸还原酶（NR）活力,氮素营养和氨基酸（AA）含量,分析了它们与生长的关系。结果表明,长白落叶松所含氮素营养以NH4－N为主,NO3－N和NO2－N含量很少。不同种源长白落叶松针叶中NH4－N和T－N（NH4－N＋N03－N＋NO2－N）含量与其NR活力呈极显著正相关。生长季初,长白落叶松针叶中氮素营养含量低,与生长呈正相关;生长季中后期,针叶中氮素营养含量高,与生长无相关性。不同种源长白落叶松针叶中AA含量的季节变化与氮素营养相似,但与生长无相关性。生长季初针叶中NR活力与不同种源长白落叶松生长呈显著正相关。初步研究表明,生长李初针叶中NR活力,NH4－N和T－N含量可作为生理生化指标用于长白落叶松种源早期选择。     

兴安、长白及华北落叶松RAPD 分子标记的物种特异性鉴定

以中国北方地区主要乡土落叶松树种兴安落叶松(Larix gmelini)、长白落叶松(L. olgensis)和华北落叶松(L. principisrupprechtii)的针叶及种子胚乳为研究材料, 采用RAPD分子标记技术对3种落叶松进行不同物种的种间鉴别。结果表明, 通过引物筛选得到了4个可以鉴别3种落叶松的RAPD引物, 其中有2个引物在落叶松针叶和种子胚乳基因组中都扩增出相同的条带。引物OPB-11在兴安和长白落叶松基因组DNA中1 500 bp处扩增出特异条带, 而在华北落叶松中没有; 引物OPX-14在兴安落叶松基因组DNA中1 200 bp处扩增出特异条带, 而在长白落叶松中没有; 还有2个引物可分别作为3种落叶松苗木和种子鉴别的辅助标记。本研究从分子水平上为落叶松的种间鉴别提供了新的鉴定方法。     

兴安、长白及华北落叶松RAPD分子标记的物种特异性鉴定

以中国北方地区主要乡土落叶松树种兴安落叶松（Larix gmelini）、长白落叶松（L.olgensis）和华北落叶松（L．principis—rupprechtii）的针叶及种子胚乳为研究材料,采用RAPD分子标记技术对3种落叶松进行不同物种的种间鉴别。结果表明,通过引物筛选得到了4个可以鉴别3种落叶松的RAPD引物,其中有2个引物在落叶松针叶和种子胚乳基因组中都扩增出相同的条带。引物OPB-11在兴安和长白落叶松基因组DNA中1500bp处扩增出特异条带,而在华北落叶松中没有：引物OPX-14在兴安落叶松基因组DNA中1200bp处扩增出特异条带,而在长白落叶松中没有：还有2个引物可分别作为3种落叶松苗木和种子鉴别的辅助标记。本研究从分子水平上为落叶松的种间鉴别提供了新的鉴定方法。     

杂种落叶松扦插生根过程中可溶性蛋白的比较分析

杂种落叶松(长白落叶松×日本落叶松)(Larix olgensis×Larix kaempferi)插穗在生根过程中,可溶性蛋白的 SDS-PAGE图谱分析推测:24KD、26KD和39KD蛋白,与根原基的发生、分化有关;28KD蛋白具有促进根原基继续发育长出不定根的作用;47KD蛋白阻遏根原基的发生或抑制不定根的生长。     

人工长白落叶松幼龄林树冠比叶重

比叶重(LMA)是构建生态系统过程模型的重要参数之一,准确预测树冠比叶重的动态变化对提高模型精度有重要意义。本研究以黑龙江省尚志市帽儿山林场人工长白山落叶松为对象,分别在生长季针叶不同发育时期对树冠内不同垂直位置的针叶比叶重进行测量,分析针叶比叶重在树冠垂直方向及针叶不同发育时期的变化规律,探讨导致其时间和空间差异的主要因子,建立长白落叶松幼龄林比叶重动态预估模型。结果表明: 比叶重在树冠垂直方向表现为随着相对着枝深度(RDINC)的增加而减小,完全展叶后比叶重在垂直方向的变化幅度明显高于展叶初期。比叶重在不同发育时期表现为随发育进程先增大后趋于稳定,该趋势随着树冠深度的增加而逐渐减弱。分别以RDINC和年度积日(DOY)为单一变量预测比叶重时,模型的调整后决定系数(R_a²)低于0.6,当同时以RDINC和DOY为自变量构建比叶重预估模型时,R_a²提高0.19,且模型检验效果良好(ME=0.54 g·m^-2, MAE=5.74 g·m^-2)。研究表明长白落叶松比叶重在树冠不同轮层和不同针叶发育期间均存在显著差异,以RDINC和DOY为自变量构建的比叶重预测模型可以很好描述长白落叶松比叶重的空间及生长季针叶发育期变化,为阐明树冠发育机理提供理论依据,为提高生态过程模型精度奠定基础。     

长白落叶松无性系抗寒性比较研究

以58个长白落叶松无性系1年生枝条为材料,对其进行低温胁迫处理,探讨不同无性系抗寒差异,结果表明：不同温度和时间的处理条件下,4个落叶松无性系电导率不同,随着低温胁迫时间的延长和温度的降低,不同无性系电导率呈上升趋势。在相同温度及时间处理条件下（-40℃、12 h）,58个长白落叶松无性系相对电导率差异达极显著水平（P<0.01）。以-40℃、12 h处理下各无性系电导率与对照的差值为指标,利用聚类法进行分析,初步筛选出9个抗寒性较强的无性系（L5、L16、L21、L23、L27、L40、L73、L78和L90）,这些无性系可作为抗寒优良无性系进行考察,本研究为落叶松抗寒良种选育提供基础。     

不同林龄、树种落叶松人工林径向生长与气候变化的关系

研究人工林径向生长与气候变化的关系对全球气候变暖背景下人工林合理经营有着重要的意义。该文对在辽东山区广泛栽培的黄花落叶松(Larix olgensis)和日本落叶松(Larix kaempferi)人工林, 运用树木年轮气候学方法建立了辽宁草河口和湾甸子林场落叶松人工林年表, 分析了落叶松径向生长对气候变化的响应以及气候条件、树种、立地条件和林分因子(林龄、密度、蓄积量等)的相对影响程度。结果发现在影响年轮-气候关系的因素中, 气象因子的潜在蒸发散(PET)的影响力最大; 林龄、密度和蓄积量同时也具有重要的影响作用。中龄落叶松人工林径向生长主要与气温呈正相关关系, 成熟落叶松人工林径向生长主要与气温呈负相关关系; 而其他因素, 如树种、立地条件等的影响作用不大。这表明在气候变暖背景下随着林龄增加, 林分会逐渐受到气温升高导致的水分亏缺的限制, 导致明显的生长下降趋势, 因而气候变暖对成熟落叶松人工林威胁更为严重, 所以要注重对成熟林的优先保护, 同时可以预测, 随着东北地区今后气候进一步变暖, 可能将逐步影响到林龄更小的林分的生长, 因此需要进一步研究如何在落叶松人工林经营中采取科学的措施来更好地应对未来气候变化。     

长白落叶松与日本落叶松的碳储量成熟龄

通过树干解析求得材积连年生长量和平均生长量,运用Vario EL Ⅲ型元素分析仪测定长白落叶松和日本落叶松不同年龄阶段碳百分含量及碳密度,并对两树种的碳储量成熟龄进行了探讨.结果表明：长白落叶松和日本落叶松的材积数量成熟龄分别为48.3年和49.3年;两树种碳密度变化趋势基本一致,长白落叶松最大值出现在30年,日本落叶松出现在35年;日本落叶松连年碳积累量大于长白落叶松,但达到最大值的年龄较晚;日本落叶松的平均碳积累量大于长白落叶松;对两树种的连年碳积累量和平均碳积累量的曲线方程进行拟合,得到长白落叶松和日本落叶松的碳储量成熟龄分别为48.7年和47.7年.     

Colour of larch heartwood and relationships to extractives and brown-rot decay resistance

Larch heartwood is appreciated for its good mechanical properties, its colour and its texture, and it is often used outdoors because of its natural durability (decay resistance). In this study the colour of larch heartwood was studied in relation to extractives and decay resistance, with the aim to estimate durability of larch heartwood from its colour. On a total of 293 trees colour in the CIE L*a*b* space (L* lightness, a* red/green axis, b* yellow/blue axis), extractives content (acetone and hot-water extractives, amount of phenolics) and the brown-rot decay resistance were determined. For calculating the relative decay resistance ( x), mass loss after inoculation for 16 weeks with two fungi [ Coniophora puteana (Schum.ex.Fr.) Karst., Poria placenta (Fr.) Cke, European standard EN 113] of larch heartwood samples was compared to Scots pine ( Pinus sylvestris L) sapwood reference samples (EN 350-1). Different species [Japanese larch ( Larix kaempferi Lamb.), Hybrid larch (Larix deciduax L. kaempferi) and European larch ( L. decidua Mill.)], provenances and age classes (38-year, >150-year) were included. Japanese larch heartwood turned out to be significantly more reddish (higher a*-values) compared to the European larch provenances. Reddishness of the hybrids was intermediate. The red hue (+a*) was strongly correlated with the amount of phenols ( r =0.84) and decay resistance ( r =0.63) and therefore suitable for prediction of both parameters. The results suggest that colour measurements of larch heartwood could be of benefit in tree breeding programs and for an optimised utilization of larch timber.     

不同光照强度下兴安落叶松对舞毒蛾幼虫生长发育及防御酶的影响

为了探讨舞毒蛾对光照环境变化引起的落叶松抗性的变化是如何响应的,分析了不同光照强度处理的兴安落叶松对舞毒蛾幼虫生长发育状况和生理指标变化的影响。研究发现,取食50%和25%光照强度下生长的落叶松后,舞毒蛾幼虫平均体重、蛹重、化蛹率和羽化率与对照组相比均显著下降（P<0.05）。幼虫体内保护酶SOD、POD、CAT和解毒酶ACP、AKP、CarE、GSTs、MFO活性与对照组(100% 光照)相比均显著降低（P<0.05）且除CAT外在50%光照强度下降低最为显著（P<0.05）;4龄和5龄幼虫取食同一光强处理的落叶松后,SOD、POD、CarE、GSTs和MFO活性5龄显著低于4龄（P<0.05）,ACP和AKP活性5龄却显著高于4龄（P<0.05）,表明昆虫在不同的发育阶段启用不同的排毒酶系。光照差异对取食人工饲料的舞毒蛾生长发育和排毒代谢酶活性均没有显著影响,说明舞毒蛾生长发育和酶活性的变化是因寄主植物抗性的影响而非光照强度的作用。本研究结果表明,采取适当的营林措施调节林分内的光照条件,可以提高落叶松的自主抗虫性,增强其抵御害虫危害的能力,有效控制害虫危害,减少化学农药的施用量。     

Response of larch species to climate changes

Global warming has changed the distributions of forests of northeastern China. Larix are very important species in this area. Predicting the potential distributions of Larix species and their responses to climate change would attract more and more attention.This paper predicted the potential distributions of three Larix species based on 'climatic-topographic' relationships by logistic regression. The results showed that L. gmelinii is predicted to retreat northwestward by 220 km by 2050 and by 270 km more by 2100; L. olgensis var. changpaiensis is predicted to retreat northwestward by 200 km by 2050 and by 190 to 300 km more by 2100; L. principis-rupprechtii is predicted to retreat northeastward by 200 km by 2050 and by 250 to 400 km more by 2100. This indicated that L. gmelinii could have its optimum latitude moved into Russia, L. olgensis var. changpaiensis could move to the Small Xing'an Mountains and L. principis-rupprechtii would move to the middle part of the Great Xing'an Mountains.     

白桦和落叶松苗木对其纯林土壤养分的生物检测

利用35年生白桦、落叶松纯林土壤分别栽植白桦、落叶松1年生苗木,通过对苗木生长、生物量、叶片养分及土壤养分变化的研究,对两种土壤的肥力进行评价,探讨白桦、落叶松混交林种间互作机制.结果表明:白桦纯林土壤全氮、碱解氮含量显著高于落叶松纯林土壤(P<0.05),落叶松纯林土壤全磷、全钾、速效磷含量显著高于白桦纯林土壤(P<0.05).盆栽第1年,白桦纯林土壤上的白桦苗木苗高、地径和生物量分别比在落叶松纯林土壤上增加69%、52%和65%(P<0.05),落叶松苗木分别增加12%、8%和37%(P>0.05).以白桦纯林土壤为基质时,白桦、落叶松苗木叶片氮浓度高于落叶松纯林土壤,而磷浓度低于落叶松纯林土壤.白桦凋落量大且分解速度快,落叶松对土壤磷具有活化作用,导致白桦纯林土壤氮素有效性较高,落叶松纯林土壤有效磷含量较高.推测两树种混交时,土壤氮、磷的互补作用可能对林分产量产生有益的作用.     

Dynamics of structure and biological productivity of post-fire larch forests in the Northern Mongolia

Peculiarities of forming and growth of post-fire larch (Larix sibirica Ledeb.) forests at the southern range of their distribution in the Northern Mongolia were studied. Regularities of the stand structure and dynamics of biological productivity are analyzed and discussed in the paper. It has been proved that the structure of the organic mass of the post-fire herb-carex type larch tree stands at the southern border of forest vegetation distribution is closely related to their biometric indices as-age, density, and productivity. In comparison with tree stands from the other areas of forest vegetation, the total phytomass stock of larch phytocenoses appropriately increases by zonal gradient from forest-tundra border to the Transbaikalian southern taiga and northern regions of Mongolia. Regeneration of forest cover by edificator and formation of the original larch coenopopulation is a positive trend from an environmental point of view, because the Siberian larch in young and middle-age period in the area has a sufficiently high growth energy and rate of phytomass production, with more than twice exceeding mature forests in fixed carbon of the atmosphere and has positive values of carbon balance and total destruction of organic matter by the “input-output” parameters.     

Nitrogen cycling and dynamic analysis of man made larch forest ecosystem

Nitrogen cycling process and dynamic change over 2 years were studied in 21-year-old planted dahurian larch (Lurix gmelinii (Rupr) Rupr.) in the eastern part of northern China. N concentrations in the plants varied by tissue, age, position in tree and season. In the aboveground components the N concentration was highest in foliage, followed by live branches, bolebark and bolewood. The organic N concentrations in undergrowth and herbs were higher than that in larch tree. The total amount of N accumulated in the larch ecosystem was 13167 kg.ha^-1, in which the percentages of N storage in soil, living plants, dead standing and litter were 94.7%, 2.8%, 0.1% and 2.4%, respectively. The uptake of N by vegetation was 56 kg.ha^-1.y^-1, in which the retention and return were 24 kg.ha^-1.y^-1 and 32 kg. ha^-1 y^-1, respectively. Precipitation provided 13 kg.ha^-1.y^-1 of N, while N loss via runoff was 4 kg.ha^-1.y^-1 and therefore, the net gain of N by ecosystem was 9 kg.ha^-1.y^-1.The simulation of N dynamic change showed that an increase in the age of stand was accompanied by a concomitant increase in N storage in the forest floor, whereas N flux from forest floor organic matter into soil would decrease, and consequently, growth rate of larch stands would reduce owing to the inadequacy of available N in the soil. The prediction indicated that the degradation of soil fertility in larch plantation might occur under continuous cropping. The study implied that release rate of N from litter into soil was the key factor in determining and regulating N cycling in larch plantation.The understory minor vegetation in the larch stand plays an important role in speeding up N cycling. The timely thinning is needed to improve growth and development of shrubs and herbs, and to avert the potential soil degradation. The mixed stand of larch with either a certain proportion of broad-leaved or a moderately well developed understory vegetation should be encouraged.     

Impacts of fire severity and post-fire reforestation on carbon pools in boreal larch forests in Northeast China

Aims Boreal larch (Larix gmelinii) forests in Northeast China have been widely disturbed since the 1987 conflagration; however, its long-term effects on the forest carbon (C) cycling have not been explored. The objective of this study thus was to quantify the effects of fire severity and post-fire reforestation on C pools and the changes of these forests.Methods Sixteen permanent plots have been set in two types of larch stands (L. gmelinii -grass, LG; and L. gmelinii-Rhododendron dahurica, LR) with three levels of fire severity (unburned, low-severity and high-severity but replanted), at 1987 burned sites in Daxing'anling, northeastern China, to repeatedly measure ecosystem C pools in 1998 and 2014. C components were partitioned into vegetation (foliage, branch, stem and roots), soil and detritus (standing and fallen woody debris and litter). The fire effects on post-fire C dynamics were examined by comparing the differences of C pools and changes between the two field investigations caused by fire severity.Important findings During the study period, unburned mature stands were C sinks (105g C m ?2 year^-1 for LG, and 190g C m ?2 year^-1 for LR), whereas the low-severity stands were C-neutral (?4 and 15g C m ?2 year^-1 for LG and LR, respectively). The high-severity burned but reforested stands were C sinks, among which, however, magnitudes (88 and 16g C m ?2 year^-1 for LG and LR, respectively) were smaller than those of the two unburned stands. Detritus C pools decreased significantly (with a loss ranging from 26 to 38g C m ?2 year^-1) in the burned stands during recent restoration. Soil organic C pools increased slightly in the unmanaged stands (unburned and low-severity, with accumulation rates ranging from 4 to 35g C m ?2 year^-1), but decreased for the high-severity replanted stands (loss rates of 28 and 36g C m ?2 year^-1 for LG and LR, respectively). These results indicate that fire severity has a dynamic post-fire effect on both C pools and distributions of the boreal larch forests, and that effective reforestation practice accelerates forest C sequestration.