氮磷添加对内蒙古温带草地地上生物量的影响

干旱半干旱地区植物生长不仅受到水分的限制,同时也受到养分的限制.为进一步明确养分在多大程度上能促进半干旱区植物的生长,本研究在内蒙古温带草原开展了连续2年的养分添加控制试验,设置10和40 g·m^-2·a^-1的N添加水平以及10 g·m^-2·a^-1的P添加水平.结果表明： N添加能够显著促进植物生长,10和40 g N·m^-2·a^-1处理地上生物量在2012年分别较对照增加50.8%和65.9%,2013年增加71.6%和93.3%,2个N添加处理之间地上生物量无显著差异.与单独10和40 g·m^-2·a^-1的N添加处理相比,P添加后地上生物量在2012年分别提高98.4%和186.8%,2013年分别提高111.7%和141.4%.N添加普遍提高了3种植物功能群(禾本科、菊科和其他科)的地上生物量,但较对照差异不显著,而N、P同时添加显著提高了菊科植物的地上生物量.养分添加使植被盖度升高,从而改善了表层土壤的水分条件,这可能是N、P添加促进植物生长和提高降水利用效率的重要机制之一.     

干旱条件对鬼针草和醉鱼草种间相互作用及生长的影响

为研究干旱对入侵植物鬼针草(Bidens pilosa)和本地种醉鱼草(Buddleja lindleyana)种间关系及生长的影响,试验设置2种栽植方式:种间组(2种植物各1株栽植于1个花盆)和挡板组(在种间组中添加挡板分离2株植物),分别进行水分处理,包括对照组(90%～100%饱和含水量)和干旱组(30%～35%的饱和含水量),测量植物形态、生物量,计算植物各特征的相对生长率和相对邻株效应指数。结果表明:(1)干旱条件下鬼针草根、茎、叶和总生物量的种间关系均为促进;而水分条件改变醉鱼草的种间关系,干旱处理为竞争,对照处理下为促进。(2)干旱下种间作用增加了鬼针草根、茎、叶生物量的相对生长率和根冠比,降低了醉鱼草比叶面积和根冠比(P<0.05)。(3)种间作用显著增加2个物种根长、根表面积和根尖数的生长(P<0.05)。研究表明,种间相互作用能提高干旱下鬼针草的生长表现,使其在干旱生境中具有优势,可能是鬼针草的入侵策略。     

长期氮添加对内蒙古典型草原植物地上、地下生物量及根冠比的影响

氮沉降的增加显著改变了生态系统功能和过程。大量研究表明,氮添加会提高生态系统生产力,然而,大部分研究集中在地上生物量,对地下生物量以及根冠比的研究较少。我们在内蒙古典型草原进行了7年的氮添加实验,设置6个氮添加水平,分别是N0(0)、N1(5.6 g N·m-2)、N2(11.2 g N·m-2)、N3(22.4 g N·m-2)、N4(39.2 g N·m-2)、N5(56 g N·m-2),测定植物地上和不同土层(0～10、10～30、30～50和50～100 cm)地下生物量并计算根冠比,研究不同氮添加水平对植物地上、地下生物量和根冠比的影响。结果表明:(1)与对照相比,低氮添加水平(N1和N2)未显著增加地上生物量,高氮添加水平(N3～N5)显著增加了地上生物量(96%～117%),且各高氮添加水平间差异不显著;(2)不同氮添加水平下,植物地上生物量的氮响应效率(NRE)均大于0且呈下降趋势,相邻氮添加水平的NRE差值(ΔNRE)仅在N3水平下显著增加,说明地上生物量在N3水平下达到饱和;(3)氮添加未显著改变不同土层以及整个土壤剖面上的地下生物量,且对各土层地下生物量的占比情况无显著影响;(4)与对照相比,N1和N2未显著改变植物的根冠比,N3～N5显著降低了植物根冠比。综上所述,氮沉降增加会提高植物地上生物量,对不同土层根系的影响比较复杂,需要更多的研究来明确地下生产力对氮沉降增加的响应规律。     

施氮对干旱胁迫下毛竹幼苗生物量和根系形态的影响

为探讨氮素添加对水分胁迫下毛竹幼苗地上生物量及地下根系形态的调控作用,选取1年生毛竹实生苗为材料,采用水分和施氮双因素完全随机区组设计,以田间持水量的80%～85%作为水分对照(CK)、50%～55%为中度干旱(MD)、30%～35%为重度干旱(HD)设置3个水分水平,氮处理分未施氮(N0,0 mg N·kg^-1)和施氮(N1,100 mg N·kg^-1)2个水平,通过盆栽试验,测定毛竹实生苗根系形态特征及各器官生物量。结果显示：施氮显著增加了同一水分下毛竹幼苗叶、根及整株生物量,其中,N1MD和N1HD分别较N0MD和N0HD地上生物量增加15.6%、11.9%,总生物量分别增加36.7%、25.0%(P<0.05);施氮降低了相同水分处理下毛竹的比根长、茎叶比,显著促进了中度和重度干旱下根冠比的增加(P<0.05);水分胁迫下,除根生物量比显著增加外,茎、叶生物量比均随氮素添加呈减小的趋势;施氮对毛竹幼苗根系形态特征(根长、根表面积、根体积)具有不同程度的促进作用;施氮对中度干旱下毛竹幼苗干物质积累的缓解作用比重度干旱大,但在...     

陆生植物生物量分配对模拟氮沉降响应的Meta分析

分析了陆生植物地上、地下各组织中生物量分配对氮沉降的响应,为研究大气氮沉降背景下陆地生态系统的碳、氮循环过程及植物生物量分配、立木收获、定向培育等相关研究和实践提供参考依据。共收集整理了国内外63篇论文的原始数据资料进行Meta分析(Meta-analysis),用以定量评估氮沉降对植物生物量分配的影响,并通过亚组分析进一步探讨了不同生态系统类型、植物种类、氮肥形式、施氮水平和持续时间对生物量分配的影响。结果表明,总体来看施氮会显著促进植物地上部分生物量分配,植物叶生物量和茎生物量在施氮条件下均显著增加;然而地下生物量所受促进作用要低于地上部分,表现为植物细根生物量和粗根生物量在氮输入下并没有显著变化;植物根冠比在氮沉降下显著降低;叶重比、茎重比和根重比在氮沉降下没有显著变化。此外,亚组分析结果表明生态系统类型和植物类型会显著影响植物总生物量和根冠比对氮沉降的响应,草本植物在氮沉降下的生物量累积明显优于木本,这说明短期氮沉降可能会增加草本的覆盖面积;施肥形式对根冠比的影响存在明显差异,相比于尿素,硝酸铵对植物根冠比的作用更显著;不同施氮水平显著影响地上生物量分配,中氮水平(本研究为60—120 kg hm-2a-1)促进作用最大,高氮水平(本研究为≥120 kg hm-2a-1)促进作用明显减弱,这与总生物量的变化一致,表明过高的氮沉降量将抑制植物生长;氮沉降处理时间长短对植物地上生物量的影响也存在显著差异,当施氮时间高于3年,氮沉降对地上生物量的促进作用几乎消失。总之,短期氮沉降会使植物分配更多生物量给地上部分,且氮沉降对草本植物生物量的累积作用明显优于木本,这些发现可为未来大气氮沉降背景下植物地上、地下部分碳存储、植物群落结构、植被动态等相关研究提供科学依据。     

两种一年生草本对喀斯特土壤厚度及供水的生长响应

以一年生草本苍耳(Xanthium sibiricum Patrin ex Widder)和鬼针草(Bidens pilosa L.)为材料,模拟喀斯特地区不同土壤厚度以及不同水分处理,研究了两者的形态及生物量积累与分配特征。结果显示:资源减少(供水、土壤厚度及供水+土壤厚度) 3种处理均抑制了植物的生长,植物生物量积累显著减少,对根和叶的投资比均未下降,而对茎的投资比显著下降;供水量减少对两者的根冠比、根长及比叶面积均无显著影响,而在土壤厚度下降及供水量与土壤厚度同时下降时,根冠比、根长与比叶面积均显著增加。研究结果表明,土壤厚度降低对2种草本植物的负面影响大于供水降低的影响。资源轻度降低时,植物以牺牲茎投资为代价保持对根和叶的稳定投资,从而保证对水分的吸收和光合合成;资源严重降低时,植物通过增加比根长等地下系统的投资以对抗地下环境压力的加剧。     

接种菌根菌剂对科尔沁沙地4种造林幼苗生长特性的影响

菌根真菌在干旱半干旱地区退化生态系统植被恢复与重建中具有重要作用。以科尔沁沙地乡土树种五角枫(Acer mono)和榆树(Ulmus pumila)及主要造林树种白蜡(Fraxinus chinensis)和樟子松(Pinus sylvestris var.mongolica)为供试植物,通过科尔沁沙地菌根化造林试验,分析4种造林幼苗接种菌根菌剂后幼苗的菌根侵染率、形态学指标、生物量积累和分配以及根际土壤养分在一个生长季内的变化。结果表明:两种菌根菌剂能较好地侵染幼苗,接种菌根菌剂显著提高了4种幼苗的株高、基径、顶枝长和生物量;显著影响幼苗的生物量分配,接种内生菌剂的白蜡根冠比显著减小,接种外生菌剂的樟子松根冠比显著增加;相关分析表明,菌根侵染率与白蜡幼苗株高和顶枝长呈显著正相关,与樟子松幼苗输导根质量、地上生物量和总生物量呈显著正相关;接种菌根菌剂还能提高根际土壤有机质和全氮含量,改良幼苗生长基质,但差异不显著。4种造林幼苗均表现出了显著的菌根效应,提示菌根菌剂在干旱贫瘠地区造林中具有很强的应用价值。     

基于容器分区处理探究黑麦草生长对喀斯特不同土壤生境和水分的响应

为了探究不同水分条件下喀斯特地区分布不均、厚薄不一土壤小生境对禾本科草本植物生长的影响,用3种不同深度的容器(对照深度CK,深土D和浅土S)两两组合为6种复合容器(CK-CK、CK-S、CK-D、D-D、S-D和S-S)以实现容器分区,研究了黑麦草的根系生长、生物量积累及其分配特征。结果表明:1)在水分充足(W_0)条件下,组合了浅土容器和深土容器的处理中,黑麦草的根系生长(根长、根直径、根表面积和根生物量)均低于对照容器(CK-CK),且有浅土容器的组合处理(S-S,S-D,CK-S)受抑制程度大于有深土容器的组合处理(CK-D,D-D);当水分含量降低后,即中水(W_1)和低水(W_2)条件下,有深土容器的组合[D-D和(或)CK-D]根系生长与对照相比显著增加,而有浅土容器的组合[S-S和(或)CK-S]根系生长与对照相比显著降低。2)对比同一处理不同容器分区中黑麦草生长指标发现,在水分充足情况下,深土容器和浅土容器均会抑制植物生长,而当水分减少,S区根系生长被严重抑制,但D区根系增长优势明显。3)水分充足条件下,根冠比未受到显著影响;当水分降低时,组合了深土容器的处理根冠比均有升高的趋势,组合了浅土容器的处理根冠比有降低趋势。由此可见,不同土壤生境带来的物理空间限制会影响植物根系生长和生物量积累与分配,但水分的减少会改变根系生长及生物量积累对不同土壤生境的响应:在水分充足时,土壤物理空间是影响根系生长和生物量积累与分配的主要因子,黑麦草主要发展浅层根系。而当水分减少时,黑麦草根系在浅层土壤中无法获取供给生长代谢活动的足量水分,更倾向于将有限的有机物分配给根,通过根系伸长、表面积和体积增大、直径增粗等策略加强水分吸收,从而增强对干旱的抗逆性,提高对土壤和水分异质性的适应。     

环青海湖地区天然草地和退耕恢复草地植物群落生物量对氮、磷添加的响应

植物群落生物量反映了植被的初级生产能力, 是陆地生态系统碳(C)输入的最主要来源, 往往受到自然界中氮(N)、磷(P)元素供应的限制。该试验以青藏高原环青海湖地区的高寒草原为研究对象, 探讨了天然草地和退耕恢复草地植被群落生物量对N (10 g·m^-2)、P (5 g·m^-2)养分添加的响应。N、P添加显著增加了天然草地禾草的生物量, 进而促使地上总生物量显著提高。退耕恢复草地禾草和杂类草的生物量对N添加均有一致的正响应, 从而促使地上总生物量显著增加174%, 群落地上和地下总生物量显著增加34%; 而P添加对恢复草地生物量各项参数均无显著影响。回归分析显示: 天然草地植物群落地上生物量随土壤中NO₃^--N含量的增加而增加(p < 0.05), 退耕恢复草地植被地上、地下和总生物量均与土壤NO₃^--N含量显著正相关(p < 0.01), 说明环湖地区高寒草原植物生长主要受N供应的限制, P的限制作用随土地利用方式的转变和群落演替阶段的不同而变化; 相比天然草地, 恢复草地在现阶段植被初级生产力受N的限制作用更强烈, 土壤中可利用N含量是限制其植被自然恢复和重建的关键因子。     

本地和外来草本物种对水分条件时间异质性的可塑响应

极端气候导致的干旱和水淹事件频发，影响了外来植物和本地植物的生长。为了解外来种和本地种植物对干旱和水淹事件发生顺序的响应，探讨草本植物适应水分时间异质性的策略，该文以美国蒙大拿州西部4种本地植物和4种外来植物为研究对象，将所有植物分别进行持续湿润(对照，CK)、水淹-干旱(I-D)和干旱-水淹(D-I)处理，并观测一系列形态和生物量特征的变化。结果表明：(1)与CK相比，D-I和I-D处理均显著降低了外来种的总生物量(P<0.05)。(2)D-I显著降低了本地种早期总生物量、后期地下生物量和根冠比，但显著提高了其后期的相对生长(P<0.05)。(3)D-I处理显著降低了所有植物的地下-地上生物量关系的异速指数，外来种异速指数显著高于本地种(P<0.05)。综上认为，极端事件(水淹和干旱)的发生顺序能改变外来植物和本地植物的生物量分配，早期干旱比后期干旱更容易减少植物生物量的积累，但能促进本地种后期的生长；本地种在环境胁迫下不被降低的总生物量表现说明维持表型稳定的能力较强；D-I处理下本地种和外来种地上和地下生物量关系的分配方式不同。     

不同水分处理下双接种丛枝菌根真菌和根瘤菌对疏叶骆驼刺生长及氮素转移的影响

以疏叶骆驼刺为研究对象,设定3个水分梯度正常水分（土壤相对含水量（70±5）%）、干旱胁迫（土壤相对含水量（20±5）%）和复水处理（干旱胁迫60天后恢复至正常水分）与四组接种处理（单接种丛枝菌根真菌（AMF）、单接种根瘤菌、双接种AMF+根瘤菌和不接种）,分析不同水分条件下双接种丛枝菌根真菌和根瘤菌对疏叶骆驼刺的生长以及供、受体疏叶骆驼刺之间氮素转移的影响。结果表明,正常水分处理时,双接种疏叶骆驼刺的AMF侵染率、地上生物量、地下生物量、总生物量以及氮含量均要高于单接种处理;根瘤数量、最大荧光（Fm）、初始荧光（Fo）、最大光化学效率（Fv/Fm）与单接种处理之间无差异;在遭遇干旱胁迫时,双接种疏叶骆驼刺的AMF侵染率、总生物量、Fv/Fm均小于单接种处理;地上生物量、地下生物量、根瘤数、Fm、Fo以及氮含量与单接种之间无差异。复水后,双接种疏叶骆驼刺的地上生物量、地下生物量、总生物量、根瘤数均优于单接种;AMF侵染率、氮含量低于单接种;Fm、Fo、Fv/Fm均与单接种之间无差异。在氮素转移方面,正常水分时,双接种与单接种的氮素转移率无差异,在遭遇干旱胁迫时,双接种疏叶骆驼刺的氮素转移率显著降低,即使复水后,仍得不到恢复。可见,与单接种AMF或单接种根瘤菌相比,双接种AMF和根瘤菌在正常水分时更具有优势,干旱胁迫会导致AMF和根瘤菌协同促生优势的减弱,复水后双接种疏叶骆驼刺能及早的对水分变化做出响应,对其生长具有一定的补偿作用,但仍不能抵消干旱胁迫所带来的损伤。丛枝菌根网络促进氮素转移一定程度上提高了疏叶骆驼刺幼苗耐旱性,但是在干旱条件下双接种疏叶骆驼刺的氮素转移率要低于单接种AMF,复水后仍得不到恢复。     

模拟喀斯特异质性小生境下三叶鬼针草地上地下协同生长对策

模拟在喀斯特异质生境下,通过随机区组实验,研究三叶鬼针草（Bidens pilosa L.）在两种土壤生境（浅而宽、深而窄）和3种水分处理（对照、减水50%、减水70%）下植物的地上和地下生长关系及生物量分配格局。结果显示：（1）两种生境中三叶鬼针草的地上生长（株高、地径、叶面积、叶生物量）与地下根系生长（根长、根表面积、根体积、根生物量）均随着施水量的减少而降低;叶面积比率随着施水量的减少而增加;根质量比在浅而宽土壤生境中呈先增后减的趋势,而在深而窄土壤生境中呈增加趋势。（2）两种生境中三叶鬼针草的地上生物量与地下根系生物量、叶面积与根长、叶面积与各层根系生长均呈显著正相关关系。但在浅而宽土壤生境中,三叶鬼针草的地上生物量与各土层根系生物量均呈显著正相关,而在深而窄土壤生境中,地上生物量仅与中上土层根系生物量呈显著正相关。研究表明三叶鬼针草在不同生境中均具有较好的地上地下协同生长对策,在增强对地下资源获取的同时也增强了对地上资源的获取。在浅而宽土壤生境中,三叶鬼针草通过协调根系的横向拓展能力与植物叶片的生长来应对快速的干旱缺水;在深而窄土壤生境中,植株能较好地协调根系向下拓展能力与地上叶面积的生长,更好地利用土壤深层的水分资源。     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Water and nitrogen addition differentially impact plant competition in a native rough fescue grassland

We examined how water and nitrogen addition and water–nitrogen interactions affect root and shoot competition intensity and competition–productivity relationships in a native rough fescue grassland in central Alberta, Canada. Water and nitrogen were added in a factorial design to plots and root exclusion tubes and netting were used to isolate root and shoot competition on two focal species (Artemisia frigida and Chenopodium leptophyllum). Both water and nitrogen were limiting to plant growth, and focal plant survival rates increased with nitrogen but not water addition. Relative allocation to root biomass increased with water addition. Competition was almost entirely belowground, with focal plants larger when released from root but not shoot competition. There were no significant relationships between productivity and root, shoot, or total competition intensity, likely because in this system shoot biomass was too low to cause strong shoot competition and root biomass was above the levels at which root competition saturates. Water addition had few effects on the intensity of root competition suggesting that root competition intensity is invariant along soil moisture gradients. Contrary to general expectation, the strength of root competition increased with nitrogen addition demonstrating that the relationship between root competition intensity and nitrogen is more complex than a simple monotonic decline as nitrogen increases. Finally, there were few interactions between nitrogen and water affecting competition. Together these results indicate that the mechanisms of competition for water and nitrogen likely differ.     

Above- and belowground responses to nitrogen addition in a Chihuahuan Desert grassland

Increased available soil nitrogen can increase biomass, lower species richness, alter soil chemistry and modify community structure in herbaceous ecosystems worldwide. Although increased nitrogen availability typically increases aboveground production and decreases species richness in mesic systems, the impacts of nitrogen additions on semiarid ecosystems remain unclear. To determine how a semiarid grassland responds to increased nitrogen availability, we examined plant community structure and above- and belowground net primary production in response to long-term nitrogen addition in a desert grassland in central New Mexico, USA. Plots were fertilized annually (10 g N m⁻²) since 1995 and NPP measured from 2004 to 2009. Differences in aboveground NPP between fertilized and control treatments occurred in 2004 following a prescribed fire and in 2006 when precipitation was double the long-term average during the summer monsoon. Presumably, nitrogen only became limiting once drought stress was alleviated. Belowground NPP was also related to precipitation, and greatest root growth occurred the year following the wettest summer, decreasing gradually thereafter. Belowground production was unrelated to aboveground production within years and unrelated to nitrogen enrichment. Species richness changed between years in response to seasonal precipitation variability, but was not altered by nitrogen addition. Community structure did respond to nitrogen fertilization primarily through increased abundance of two dominant perennial grasses. These results were contrary to most nitrogen addition studies that find increased biomass and decreased species richness with nitrogen fertilization. Therefore, factors other than nitrogen deposition, such as fire or drought, may play a stronger role in shaping semiarid grassland communities than soil fertility.     

Above- and belowground response of Populus grandidentata to elevated atmospheric CO2 and soil N availability

Soil N availability may play an important role in regulating the long-term responses of plants to rising atmospheric CO₂ partial pressure. To further examine the linkage between above- and belowground C and N cycles at elevated CO₂, we grew clonally propagated cuttings of Populus grandidentata in the field at ambient and twice ambient CO₂ in open bottom root boxes filled with organic matter poor native soil. Nitrogen was added to all root boxes at a rate equivalent to net N mineralization in local dry oak forests. Nitrogen added during August was enriched with ¹⁵N to trace the flux of N within the plant-soil system. Above-and belowground growth, CO₂ assimilation, and leaf N content were measured non-destructively over 142 d. After final destructive harvest, roots, stems, and leaves were analyzed for total N and ¹⁵N. There was no CO₂ treatment effect on leaf area, root length, or net assimilation prior to the completion of N addition. Following the N addition, leaf N content increased in both CO₂ treatments, but net assimilation showed a sustained increase only in elevated CO₂ grown plants. Root relative extension rate was greater at elevated CO₂, both before and after the N addition. Although final root biomass was greater at elevated CO₂, there was no CO₂ effect on plant N uptake or allocation. While low soil N availability severely inhibited CO₂ responses, high CO₂ grown plants were more responsive to N. This differential behavior must be considered in light of the temporal and spatial heterogeneity of soil resources, particularly N which often limits plant growth in temperate forests.     

Effects of water supply on root traits and biological yield of Durum (Triticum durum Desf.) and Khorasan (Triticum turanicum Jakubz) wheat

The ability of a plant to change its root characteristics to increase the acquisition of soil water is an important adaptation mechanism to water limitation. In this regard, a field study was carried out in the Pannonian region of eastern Austria with two tetraploid wheat genotypes, i.e. Durum (Triticum durum Desf.) and Khorasan (Triticum turanicum Jakubz), during a comparatively wetter and drier year, i.e. 2008 and 2009, respectively. The genotypes showed significant differences in average root diameter and fine root length. All root traits varied with soil depth. The highest root length density and root biomass were observed with Khorasan wheat in 0–10 cm soil depth. Durum wheat showed a stronger response in fine roots to water availability and produced more fine roots in the moist year. Electric root capacitance was higher with Khorasan wheat. Durum showed higher biological yield stability across years with different precipitation with respect to above- and belowground biomass. It produced more leaf area under humid conditions. Khorasan allocated more assimilates to belowground organs in dry conditions, but without positive effect on aboveground biomass.     

Above- and belowground biomass allocation and its regulation by plant density in six common grassland species in China

Above- and belowground biomass allocation is an essential plant functional trait that reflects plant survival strategies and affects belowground carbon pool estimation in grasslands. However, due to the difficulty of distinguishing living and dead roots, estimation of biomass allocation from field-based studies currently show large uncertainties. In addition, the dependence of biomass allocation on plant species, functional type as well as plant density remains poorly addressed. Here, we conducted greenhouse manipulation experiments to study above- and belowground biomass allocation and its density regulation for six common grassland species with different functional types (i.e., C₃ vs C₄; annuals vs perennials) from temperate China. To explore the density regulation on the biomass allocation, we used five density levels: 25, 100, 225, 400, and 625 plant m^?2. We found that mean root to shoot ratio (R/S) values ranged from 0.04 to 0.92 across the six species, much lower than those obtained in previous field studies. We also found much lower R/S values in annuals than in perennials (C. glaucum and S. viridis vs C. squarrosa, L. chinensis, M. sativa and S. grandis) and in C₄ plants than in C₃ plants (C. squarrosa vs L. chinensis, M. sativa and S. grandis). In addition to S. grandis, plant density had significant effects on the shoot and root biomass fraction and R/S for the other five species. Plant density also affected the allometric relationships between above- and belowground biomass significantly. Our results suggest that R/S values obtained from field investigations may be severely overestimated and that R/S values vary largely across species with different functional types. Our findings provide novel insights into approximating the difficult-to-measure belowground living biomass in grasslands, and highlight that species composition and intraspecific competition will regulate belowground carbon estimation.

     

Resource competition and suppression of plants colonizing early successional old fields

Early colonizing annual plants are rapidly suppressed in secondary succession on fertile midwestern old fields, while later colonizing perennials persist. Differences in competitive ability for above- and belowground resources may be partly responsible for differences in species persistence during succession, as both light and nutrient availability may change rapidly. We found that, although both above- and belowground competition suppress growth of colonizing plants, belowground competition was the dominant factor in the suppression of the annual Ambrosia artemisiifolia in 2nd-year-old fields near the W.K. Kellogg Biological Station in southwestern Michigan. Despite an ability to persist in later successional fields, seedling transplants of the perennial Achillea millefolium were also suppressed by above- and belowground competition, with belowground competition having the strongest effect. As in many old fields, nitrogen availability is the primary factor limiting plant productivity. There was no clear difference between the species in ability to compete for ¹⁵N from an enriched patch, although there was an indication of greater precision of foraging by Achillea. Life history differences between these species and consequent differences in the phenology of root growth relative to other old-field plants are likely to play a large role in the persistence of Achillea in successional fields where Ambrosia is suppressed. Received: 8 January 1998 / Accepted: 16 September 1998