扎龙湿地异质生境芦苇种群根茎动态及年龄结构

无性系植物根茎不仅具有营养繁殖和扩展种群的功能,也是无性系家族中芽和分株生理整合的通道。采用单位土体挖掘取样,对扎龙湿地单优种群落芦苇种群根茎进行调查,比较了湿生生境和水生生境根茎长度、生物量和干物质贮量的季节变化。结果表明:芦苇种群根茎长度、生物量和干物质贮量均以湿生生境显著大于水生生境;8月前,根茎长度缓慢增加,8月后,根茎长度迅速增加,以3龄最长,6龄最短;根茎生物量、干物质贮量先下降后上升,8月最低,至10月休眠期均已超过营养生长初期,生物量以3龄最大,1龄最小,干物质贮量以5龄最高,1龄最低;不同龄级根茎长度及年龄谱与月份间呈线性相关(R~20.81,P0.05),不同龄级根茎生物量与月份间呈二次曲线关系(R~20.98,P0.05),生物量年龄谱与月份间呈线性相关(R~20.80,P0.05),5个生育期根茎干物质贮量与龄级之间呈二次曲线关系(R~20.86,P0.05)。芦苇种群根茎长度、生物量、干物质贮量有着相同的季节变化规律,不同龄级根茎的寿命与养分的输出消耗和养分的再输入补偿密切相关。整个生育期内,生境异质性对芦苇种群根茎动态及年龄结构的影响均存在,且引起的差异也相对较稳定。     

扎龙湿地不同生境芦苇种群根茎构件的年龄结构

芦苇是典型的长根茎型克隆植物,天然种群主要依靠根茎的营养繁殖进行更新。采用单位土体挖掘取样,按实际生活年限划分根茎龄级的方法,对扎龙湿地芦苇种群不同龄级根茎进行调查。结果表明,6月份,4个生境芦苇种群根茎构件均由2—6a的5个龄级组成,7—10月份均由1—6a的6个龄级组成。6—10月份,1a根茎长度比率逐渐增加;除盐碱生境略有增加外,其他3个生境2a、3a比率均小幅减少;4—6a比率逐渐减小,根茎长度比率均以3a最大,依次是2a,4a,1a,5a,最高的6a最小。1—3a根茎生物量比率逐渐增加;4—6a比率逐渐减小,均以3a根茎生物量比率最大,依次是4a,2a,5a,6a,以最低1a最小。不同龄级根茎长度和生物量比率与返青后实际生长时间之间均较好地符合直线函数关系(R²>0.91,P<0.01;R²>0.81,P<0.05),6—9月份,根茎长度呈衰退型年龄结构,10月份又转为稳定型结构,整个生长季根茎生物量均为衰退型结构。不同龄级根茎构件在芦苇种群中的地位和作用不同,根茎构件的年龄结构蕴含着种群调节的重要信息。     

扎龙湿地不同生境芦苇种群根茎数量特征及动态

采用单位土体取样,计测长度和生物量的调查与统计方法,对扎龙湿地保护区4个生境芦苇种群根茎数量特征进行比较分析。结果表明,芦苇5月10日左右返青后进入营养生长期,根茎长度6—8月份缓慢增加,8—10月份显著增加,后期是前期的3.5—10.3倍,生长季中后期是种群新根茎补充和生长的主要时期,不仅实现了种群空间扩展,并为营养繁殖储备更多繁殖芽;根茎生物量和干物质贮量6—8月份逐渐减少,8—10月份又逐渐增加,均以生长季末期的10月份最大,并均显著地(P0.05)高于其他月份,种群根茎养分的消耗主要供给根茎芽的萌发和幼株生长,根茎养分的储藏又为翌年种群的更新及扩展提供物质保障,种群对地下根茎存在明显的养分"超补偿性"贮藏现象。种群根茎长度和生物量均以湿生生境最大,依次为旱生生境、水生生境,盐碱生境最小,根茎干物质贮量以旱生生境最大,依次为湿生生境、水生生境,盐碱生境最小。种群根茎长度与返青后实际生长时间之间均较好地符合直线函数关系,种群根茎生物量和干物质贮量与生长时间之间较好地符合二次曲线函数关系,R2在0.804—0.997之间,拟合方程均达到了显著或极显著(P0.01)水平。4个生境芦苇种群在根茎长度、生物量、干物质贮量等数量特征均表现出由遗传因素控制的比较稳定的季节动态规律,在生境间的差异及其差异序位又均基本稳定,均表现出明显的土壤因子环境效应,其中土壤含水量、有机质、速效氮为正向驱动,p H、速效磷为负向驱动,土壤含水量、p H对根茎数量特征的驱动作用更突出。     

松嫩平原旱地生境芦苇种群不同龄级根茎的干物质贮藏及水溶糖含量

通过对松嫩平原旱地生境单优种群落芦苇种群不同龄级根茎的调查测定,分析了一个生长季内3个生育期芦苇种群各龄级根茎干物质贮量和水溶糖含量的季节变化.结果表明:各生育期低龄级芦苇种群根茎中干物质贮量和水溶糖含量均较少,且与高龄级的差异较大;随着生长季的进程,低龄级根茎中干物质贮量和水溶糖含量迅速增加,且与高龄级的差异逐步缩小.在整个生长季,芦苇种群各龄级根茎均具有养分消耗和再贮藏乃至超补偿性贮藏的活性,其中,幼龄级根茎的活性更大.高龄级根茎的干物质贮量和水溶糖含量有一个逐年累积的增加过程.根茎干物质贮量在龄级间和龄级内的差异均达到了极显著水平(P<0.01),并且龄级间的差异大于龄级内.水溶糖含量仅在龄级间差异达到显著水平(P<0.05).随着龄级的增加,芦苇种群根茎的干物质贮量和水溶糖含量均呈二次曲线变化.     

黑龙江扎龙湿地芦苇种群构件数量特征及其相关性

采用刈割和挖土取样,对扎龙湿地旱生、湿生、水生和盐碱生境芦苇种群分株、根茎和根茎芽进行调查。结果表明:6—10月,4个生境芦苇种群分株密度、根茎长度和根茎芽库存量、输入量、休眠量均以湿生生境最大,盐碱生境最小;根茎芽输出量均以水生生境最大,盐碱生境最小,构件间的差异及其差异序位均相对稳定,构件种群均存在明显的环境效应,构件水平上存在较大的可塑性;构件间又均表现出一定的协同进化性,其中分株密度与根茎长度、根茎芽库存量和输入量之间呈显著或极显著正相关,与根茎芽休眠量之间呈显著负相关;根茎芽库存量、输入量和输出量与根茎长度之间呈显著或极显著正相关;芦苇分株均由根茎芽萌发形成,根茎芽对分株的贡献率为100%,80%以上的根茎芽萌发形成新根茎,根茎的生长又可形成更多的根茎芽,不同构件形态结构的更替改变维持着种群的稳定和持续更新。     

松嫩平原碱化草甸旱地生境芦苇种群的芽流和芽库动态

在松嫩平原碱化草甸,旱地生境芦苇种群的根茎分布在土层约1m的不同深度,一般可生活6个年度,个别根茎可存活7～9年,乃至更长的时间.通过芦苇根茎芽调查,创建了植物种群的芽流模型.提出了采用当年1龄级根茎芽的输入率与其它龄级休眠芽库存率之和估计芦苇种群芽库贮量动态的方法.结果表明,随着生长季的进程,芦苇种群芽库输入率呈不断增加趋势,而萌发输出率呈不断减少的趋势,死亡输出率则大体保持相同的较低水平.至休眠前期的9月底,芽库输入率已为输出率的2.04倍.在松嫩平原碱化草甸旱地生境,芦苇种群各龄级根茎的休眠芽有一个稳定的萌发输出过程.定量分析结果表明,芦苇种群不同龄级根茎的休眠芽每年都有11%的比率萌发形成1龄级新根茎.1龄级根茎顶端翌年发育为分蘖株后,可为直接相连接的老龄级根茎就近输送养分,从而实现老龄级根茎芽的活力.     

东北草地异质生境芦苇种群根茎芽年龄结构及输出规律

为明确异质生境条件下芦苇种群根茎芽年龄结构及输出规律,揭示芦苇种群的营养繁殖特性,采用单位土体挖掘取样,分别计数各龄级根茎芽的调查与统计方法,对东北草甸草原草甸土和盐碱土两个生境单优群落芦苇种群根茎芽动态进行比较分析。结果表明,两个生境芦苇种群根茎芽库主要均由6个龄级组成;草甸土生境在6—10月均为增长型年龄结构;盐碱土生境6—7月份为衰退型年龄结构,8月份为稳定型年龄结构,9—10月份为增长型年龄结构。根茎芽数量1—4a普遍以草甸土生境高于盐碱土生境,5—6a普遍以盐碱土生境高于草甸土生境,各龄级根茎芽数量与月份之间均符合y=a+bx直线关系(P0.05)。随着龄级的增加,休眠芽比率呈逐渐下降趋势,而萌发芽比率则呈逐渐上升趋势,5个生育期的休眠芽比率和萌发芽比率与龄级之间均符合y=a+bx直线关系(P0.01)。各龄级根茎的休眠芽具有一个相对稳定的萌发输出过程,草甸土生境根茎休眠芽按每年11%的比率萌发输出,而盐碱土生境根茎休眠芽按每年7%的比率萌发输出。虽然芦苇种群根茎芽年龄结构及年龄谱在异质生境中存在显著差异,但却有着相同的季节变化规律,均以不断形成新根茎的芽来维持着种群的营养繁殖更新。     

荒漠绿洲区芦苇种群构件生物量与地下水埋深关系

在野外调查的基础上 ,利用 Pearson关联分析法研究了河西荒漠绿洲区不同生境中芦苇种群构件生物量与地下水埋深的关系 ,结果表明 :地下水埋深对芦苇种群的影响明显 ,主要体现在 :在湿地和地下水埋深小于 0 .5 m的生境中 ,芦苇种群表现为高的种群密度、低的种群高度、低的地上生物量和高的地下生物量 ;在地下水埋深为 1 .5 m左右的生境中 ,芦苇种群地上生物量最高 ,生长最旺盛。随着地下水埋深的增加 ,垂直根茎生物量对芦苇种群地上生物量的影响加强 ,而水平根茎生物量对芦苇种群生物量的影响呈随机型 ,这可能是造成芦苇种群呈不连续、斑块状分布的主要原因。在地下水埋深 1 .5 m左右的生境中 ,芦苇的垂直根茎对种群生物量影响十分显著 ( P<0 .0 1 ) ;在地下水埋深 1 .5～ 4.0 m的生境中 ,影响显著 ( P<0 .0 5 ) ,但在地下水埋深 4m以上的生境中影响不显著 ,这表明在河西荒漠绿洲区 ,芦苇垂直根茎的最大水力提升高度 4m左右     

扎龙湿地保护区异质生境芦苇种群分株构件的数量特征

采用单位面积取样,计数和测量的调查与统计方法,对扎龙湿地保护区4个生境单优群落芦苇种群分株构件数量特征进行比较分析。结果表明,4个生境芦苇种群从5月10日左右返青后进入营养生长期,分株高度和分株密度均持续增加到生殖生长初期的8月份,其中6-8月份差异均达到显著水平(P0.05),8-10月份差异均未达到显著水平(P0.05),芦苇进入生殖生长期后,分株便停止高度生长,地下芽的输出也不再形成分株补充现实种群;分株生物量和种群生物量均持续增加到生殖生长旺盛期的9月份,至休眠期的10月份均有所降低,各月份间的差异均达到显著水平,芦苇种群在生长季末期,具有将生产的物质分配转移到地下储藏与营养繁殖器官的形成与生长上的特性。芦苇种群分株数量特征与返青后实际生长时间之间均较好地符合对数函数关系,R~2在0.818-0.994之间,拟合方程均达到了P0.05的显著水平。4个生境芦苇种群分株密度以湿生生境最大,依次为水生生境、旱生生境,盐碱生境最小,分株高度、分株生物量和种群生物量均以水生生境最大,依次为湿生生境、旱生生境,盐碱生境最小。因此,4个生境芦苇种群分株构件数量特征均表现出基本一致的生育期节律性,同时,芦苇种群的个体生长和种群动态存在明显的环境效应,土壤含水量、pH是影响该地区芦苇分株数量特征的主要因子。     

扎龙湿地芦苇分株生态可塑性及其对土壤因子的响应

扎龙湿地的芦苇既可形成大面积的单优群落,也可形成不同群落斑块。采用大样本抽样调查与统计分析方法,对湿地内水生生境、湿生生境、旱生生境和盐碱生境芦苇种群分株高度和生物量进行比较。结果表明,6—10月份,4个生境芦苇种群分株高度及生物量均以水生生境最高,盐碱生境最低,水生生境株高为盐碱生境的1.5—2.3倍,分株生物量为2.0—5.1倍,生境间的差异性以及差异序位均相对稳定。4个生境株高生境间变异系数(19.45%—31.56%)均高于生境内变异系数(8.07%—17.61%),分株高度在生境间的可塑性更大;分株生物量中水生生境、湿生生境和盐碱生境3个生境间的变异系数(33.43%—55.61%)均低于生境内变异系数(44.85%—79.82%),分株生物量在生境内的可塑性更大。不同生境条件下芦苇种群分株,在生长和生产上均存在较大的生态可塑性,表现出明显的环境效应,其中土壤含水量是该地区芦苇分株生态可塑性变异的主要驱动因子(R0.80),为正向驱动。     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

基因型多样性对羊草种群年龄结构的影响

基因型多样性作为遗传多样性的重要组成部分,不仅在群落水平,而且在种群水平均可以发挥重要生态作用。以羊草（Leymus chinensis）为研究对象,设置5种基因型多样性梯度试验控制小区,进行基因型多样性对羊草种群年龄结构的影响研究。结果表明：随基因型多样性梯度的逐步增加,2a分蘖株生物量和数量均显著增加（P<0.05）;1-4龄级羊草根茎生物量、长度和干物质积累量多数呈极显著性增加的变化趋势（P<0.01）。在1、2、4、8和12种基因型多样性梯度时,分蘖株生物量及数量年龄结构均以1a或2a分蘖株所占比例为最高,年龄结构表现为增长型或稳定型;而羊草种群根茎生物量和长度也以2a根茎所占比例为最高,年龄结构表现为稳定型。随基因型多样性水平的增加,羊草种群分蘖株数量和生物量的年龄结构从增长型向稳定型过渡,但根茎长度和生物量的年龄结构均为稳定型。12基因型多样性水平的4龄级分蘖株和根茎的特征高于其他基因型多样性水平。因此,在不同基因型多样性水平羊草种群分蘖株和根茎具有增长型或稳定型的年龄结构,适当的基因型多样性数量显著促进了各龄级羊草分蘖株和根茎的生物量和数量特征的升高,但高度的基因型多样性水平会增加羊草种群中高龄级的分蘖株和根茎所占的比例,种群发展表现出衰退信号。     

Clonal organization of populations of Asarum europaeum and Maianthemum bifolium in contrasting woodland habitats

Populations of two rhizomatous species, Asarum europaeum (asarabacca) and Maianthemum bifolium (May lily), were examined in two, and four forest habitats respectively, in the Roztocze National Park (south-eastern Poland). May lily populations were studied in habitats: the Carpathian beechwood, upland mixed fir forest, subboreal moist mixed coniferous forest and bog-alder forest. Asarabacca was studied in two habitats: beechwood and Scots pine community (an 80-year-old plantation). In both the species studied intra- and inter-populational differences of the size of genets in terms of above- and below-ground parts of individuals as well as the biomass and area occupied were observed. In May lily populations the greatest mean number of shoots per genet was found in the fir forest (11.62±3.29), a value almost twice as great as that in the moist coniferous forest and nearly three times greater than in the bog-alder forest. Total rhizome length was also the greatest in the fir forest (351.9±98.7 cm) followed by moist coniferous forest, beechwood and alder forest habitats. In all populations of May lily a greater part of total dry weight biomass is in below-ground organs. The greatest biomass value of a genet was found in the fir forest (4.275 g), the smallest in the bog-alder forest (0.110 g). All populations differed significantly in terms of leaf area, leaf length (with the exception of fir forest and beechwood habitats where the values were the greatest), and leaf width (excluding moist coniferous and bog-alder forests which had the smallest values). In the case of asarabacca, both the mean number of ramets per genet (3.36±0.45 vs. 2.49±0.20) and total rhizome length (40.3±6.4 cm vs. 21.1±1.8 cm) were greater in the beechwood habitat than in the pine community. In the first population genets had 3–5 times greater the total biomass of those from the pine community. Only genets of the latter had proportionately more dry weight biomass in above-ground parts. It seems to be correlated with greater rhizome dieback and disintegration of genets into smaller units. Both populations were significantly different in terms of all examined parameters of leaves. Genets of both the species studied were found to have their own structure of developmental phases that often differed for shoots and rhizomes.     

Effects of rhizome age on the decomposition rate of Phragmites australis rhizomes

The change in dried rhizome samples that were left to decompose was investigated to elucidate the effects of rhizome age on the decomposition rate of Phragmites australis. Rhizomes were classified into five age categories and placed 30 cm below the soil surface of a reed stand. After 369 days of decay, new (i.e., aged less than one year) rhizomes had lost 84% of their original dry mass, compared with a loss of 41–62% for that of older rhizomes. The exponential decay rates of older rhizomes were nearly identical to that of aboveground biomass. The nitrogen (N) concentration increased to two times its original values, but the phosphorus (P) concentration remained constant after an initial loss by leaching. The carbon to nitrogen (C:N) and carbon to phosphorus ratios (C:P) leveled out at 22:1 to 38:1 and 828:1 to 1431:1, respectively, regardless of rhizome age. The results are important to understand the nutrient cycles of reed-dominant marsh ecosystems.     

Verification of a mathematical growth model ofPhragmites australis using field data from two scottish lochs

A growth model ofPhragmites australis was verified using two independent sets of published field data. The model simulates the growth pattern of a well-established, monospecific stand ofP. australis in the absence of genetic diversity and environmental stresses of mainly nutrient and water deficiency. The model formulated using first order differential equations was combined with plant phenology and comprises five subroutines in which photosynthetically active radiation, shoot, root, rhizome and new rhizome biomass are calculated. Using the model, experimental results were reproduced within reasonable limits having concordance correlation coefficients of more than 0.75 for 70% of the output parameters, which was the main objective of the study. The modelled efficiencies of PAR were 7.15% and 3.09%, as opposed to 7.7% and 2.53% in experimental estimations, for Loch of Forfar and Loch of Balgavies, respectively. Production and seasonal fluxes of dry matter ofP. australis in Scottish lochs were estimated using the modelled quantities for the 1975 growing season in g m⁻². They showed that 31% and 37% of total net photosynthate translocated to rhizomes before shoot senescence began in Loch of Forfar and Loch of Balgavies, respectively. Also in both lochs approximately 45% of total downward translocation came from accumulated shoot dry matter during senescence, while the rest came from photosynthesis before the shoots started to senesce.