克隆整合在空心莲子草抗病原真菌中的作用

入侵克隆杂草——空心莲子草在其入侵地给自然环境及社会经济造成严重危害,研究发现其野外种群很少感染病害。克隆整合作用在空心莲子草的生长发育中起着重要的作用,然而克隆整合作用对其抗病性影响的研究却很少。因此,设置实验以验证克隆整合在空心莲子草的抗病原真菌中是否起重要的作用。试验结果表明,空心莲子草的抗病原真菌能力显著强于同科植物苋菜,在接种病原真菌立枯丝核菌40 h后,苋菜叶片的侵染面积达到50%以上,而空心莲子草叶片的侵染面积不到10%;且克隆整合作用促进了空心莲子草顶端分株的地上部分的生长,茎段相连处理下,顶端的茎长、茎节数、茎的生物量、叶片数量、叶片生物量都显著高于基端,而茎段剪切处理下,顶端分株的地上部分生长并不显著大于基端分株;叶片接种病原真菌立枯丝核菌后,相连的顶端分株的总酚含量显著高于切断分株的顶端分株的含量。且二次接种病原菌后,相连分株的病原真菌侵染叶片面积不到0.1 cm~2,显著小于剪切分株的叶片侵染面积(0.8 cm~2)。结果表明,克隆整合作用不仅促进了空心莲子草相连分株的地上部分的生长,也提高了其抗病原真菌的能力。     

基于复合节间组分关系的结缕草克隆植株形态可塑性特征分析

结缕草Zoysia japonica克隆植株的基本形态学单元是复合节间。分析复合节间组分关系,可以进一步揭示其形态可塑性的细微特征和资源传输与分配的差异,同时也可以为形态可塑性成因提供合理的生理生态学解释。本文对温室内、外两种环境条件下生长的结缕草克隆植株的复合节间组分进行测定和分析,结果表明：温室内、外植株主匍匐茎的长度和生物量的波动比一级分枝大,缩短节间长度比伸长节间长度稳定,但缩短节间生物量较伸长节间生物量变化幅度大。伸长、缩短节间长度、生物量分别与复合节序数呈二次函数关系。伸长、缩短节间生物量和分株生物量在一个生长季的生长方程亦均呈二次函数关系。主匍匐茎根数量、生物量随复合节数递增基本呈递增趋势。在两种环境下,A分株根数、生物量均大于B分株根数、生物量（在结缕草匍匐茎上,着生于每个复合节的基端和梢端缩短节间上的分株分别称为A、B分株）,且差异明显。温室内植株,在分株发育形成后,伸长、缩短节间的长度、生物量分别与匍匐茎根、A分株根数、B分株根数、生物量间相关性显著,温室外的相关性格局较复杂,且总体显著性低于温室内。结缕草克隆植株基本形态结构单元的复合节上的各个组成部分之间存在的密切相关关系,是导致结缕草克隆植物形成形态可塑性的细微特征格局的主要原因。     

空心莲子草在不同生境中氮素迁移和表型可塑性差异

为探究克隆繁殖性入侵植物空心莲子草在不同生境下的生长特性,以更好地了解其整体的生存策略,利用15N同位素标定技术评估空心莲子草在陆生(Ⅰ)、湿生(Ⅱ)、水生(Ⅲ)3种不同生境相互转换中的氮素迁移效率,并对不同生境中的空心莲子草生物量分配和表型可塑性差异进行分析研究。结果表明,在不同生境中,空心莲子草氮素迁移量和各构件的生物量均存在显著差异。氮素迁移量存在显著差异(P0.05),具体生境体现为ⅡⅢⅠ;Ⅱ较Ⅲ平均增长23.32%,较Ⅰ平均增长54.49%。空心莲子草植株总生物量Ⅱ最大,Ⅰ最小。地下生物量投资与生境含水量呈显著负相关,体现为ⅠⅡⅢ,茎生物量投资比叶生物量投资平均高15.57%。在不同生境中,空心莲子草株高和茎节长度均体现为ⅡⅢⅠ,陆生、湿生、水生3种不同生境的母株处理下,Ⅱ株高分别较Ⅲ和Ⅰ增长39%、189%,52%、116%,56%、127%;茎节长度分别增长36%、150%,63%、117%,57%、94%。空心莲子草在不同生境下的氮素迁移量、各构件的生物量以及分配格局表现出的显著差异显示了其在湿生生境下具备更强的氮素迁移利用、表型可塑性和对异质环境的适应能力,揭示了克隆繁殖植物空心莲子草在湿地生境中具有更强的入侵性。该研究可为空心莲子草防控提供理论指导依据。     

不同海拔高度短穗兔耳草克隆生长及克隆繁殖特征

研究了青藏铁路和青藏公路沿线 5个不同海拔高度样地中短穗兔耳草克隆生长和克隆繁殖的特征.结果表明,不同海拔高度短穗兔耳草匍匐茎数量、匍匐茎长度、基株干重、匍匐茎干重和无性系分株干重等均存在极显著差异(P<0. 01);匍匐茎数量与海拔高度之间存在明显的负线性相关性(P=0. 046 4);匍匐茎长度、基株干重、匍匐茎干重、单位长度上匍匐茎干重、无性系分株干重和匍匐茎分株总干重 /基株干重比值均与海拔高度之间存在极显著的二次多项式相关性(P<0. 01),但匍匐茎长度、匍匐茎分株总干重 /基株干重随海拔高度的变化趋势与基株干重、匍匐茎干重、单位长度上匍匐茎干重、无性系分株干重随海拔高度的变化趋势相反.在短穗兔耳草分布的海拔范围内,其克隆繁殖和克隆生长有一个“最佳”海拔高度,远离这一高度,其克隆繁殖和克隆生长会受到一定限制. 图 6表 2参 18     

空心莲子草入侵对土壤AM真菌生物量和群落结构的影响——以湖北省典型区域为例

土壤丛枝菌根(Arbuscular mycorrhizal,AM)真菌具有维持植被稳定性和多样性的重要作用.为了解空心莲子草入侵过程中与AM真菌的相互作用,采用脂肪酸甲酯(FAMEs)和基于18S rDNA的PCR-DGGE方法对湖北咸宁、仙桃和武汉三地空心莲子草(Alternanthera philoxeroides)入侵前后土壤AM真菌的生物量和群落结构进行了比较分析.结果发现,基于FAMEs表征的AM真菌生物量在空心莲子草入侵后变化不大,仅仙桃地区显著增加(P<0.05).尽管DGGE图谱显示咸宁、仙桃和武汉三地AM真菌优势类群的群落结构均发生了明显演替,但群落间物种组成差异的多样性指数βWH在2.55-2.73之间、群落多样性指数H′在1.22-1.38之间,变化均不大.结果表明,空心莲子草入侵所形成的植物更替明显改变了AM真菌优势类群的结构,但对AM真菌的生物量和多样性影响很小.     

模拟采食下克隆整合对虉草生长及生理特征的影响

为了解克隆整合对克隆植物适应均质性采食环境胁迫的作用,以江西省鄱阳湖-乐安河段河岸带优势根状茎克隆物种虉草(Phalaris arundinacea)为对象,研究均质性采食条件下虉草的生理生态响应以及克隆整合在虉草应对模拟采食两月后发挥的作用.模拟采食(去叶)强度设置4组水平:不去叶(CK对照组)、去叶25%、去叶50%和去叶75%;克隆整合采用2种方式:茎切断和连接.结果显示,无论是否进行断茎处理,模拟采食均显著(P5%)降低虉草的叶片数、地上枝总长度、分株数和生物量,去叶处理下断茎的虉草分株相对茎连接分株具有更强生长优势.断茎和去叶处理对虉草叶绿素含量无显著影响,断茎的虉草分株在25%、50%去叶水平下叶绿素a(Chl a)、叶绿素b(Chl b)和总叶绿素(Chlt)含量均高于茎连接的分株,所有去叶处理强度下断茎的虉草分株各叶绿素指标均高于对照组.去叶显著影响了虉草分株各光合指标,断茎分株在50%、75%去叶水平上的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、胞间CO2浓度(Ci)等指标均高于茎连接分株,25%去叶处理组则相反;去叶处理的虉草分株的Pn、Tr、Ci均低于对照组.因此,采食引起虉草分株光合速率下降,使营养物质积累不足,导致生长状态下降;虉草能够在一定程度上适应均质性采食处理带来的压力,但是克隆整合不能显著改善均质性采食条件下虉草的生长.     

入侵种喜旱莲子草对土壤养分的表型可塑性反应

对外来入侵种喜旱莲子草（Alternanthera philoxeroides（Mart．）Griseb．）在不同土壤氮养分处理中的表型可塑性反应进行了研究。结果表明：土壤中氮养分水平对喜旱莲子草克隆生长有显著影响，如分枝强度、苇节长度随氮含量升高均有不同程度的增加；氮养分最高的处理中根系生物量投资最小，而茎、叶生物量投资最大，对照处理中则表现山根冠比最大。研究结果表明，在不同土壤养分条件下，喜旱莲子草不同表型结构有可能同时对环境选择作出反应，在种群生存与维持、生长、繁殖等功能方面实现种群个体各器官生物量投资的优化配置来适应多样化的环境，以增强其入侵能力，     

空心莲子草叶甲释放量对空心莲子草防控效果的研究

空心莲子草(Alternantheraphiloxeroides)是一种多年生的全球性恶性杂草,破坏入侵地生物多样性,给航运及水产养殖业带来巨大损失。利用生物防治可以有效控制空心莲子草(Alternantheraphiloxeroides)的扩散蔓延,是国内外的研究热点之一。空心莲子草叶甲(Agasicles hygrophila)是从美国引入的专食性天敌昆虫,对空心莲子草有很好的防治效果,且具有生态环保性良好、投入低、防效长等特点。天敌昆虫的释放量可显著影响防控效果,有关其最佳释放量的研究对实际应用具有重要指导意义。依托空心莲子草天敌工厂,采用环境条件控制试验,对空心莲子草的种群密度、植株形态、叶绿素含量、生物量等生物学和生理生化指标进行测定,结合对各种条件下空心莲子草叶甲种群数量的分析,研究了不同天敌释放量(0～8对·m-2)对空心莲子草的防治效果。结果表明,在初始种群密度为40 stems·m-2条件下,随着释放量的增加,空心莲子草叶甲成虫和羽化孔的数量均呈现先增加后减少的趋势,而卵、各龄幼虫、蛹的数量变化规律各不相同;空心莲子草的茎密度、生物量(湿质量、干质量)均随天敌释放量增加而逐渐降低,而节间距、茎直径等形态指标则呈现先降低后增加的趋势。研究结果显示,释放4对空心莲子草叶甲可达到最佳防治效果。在此释放量下,空心莲子草叶甲羽化孔数量和成虫数量均达到最高,空心莲子草种群密度最低,测得的节间距和茎直径最短,叶片中叶绿素含量较低,生物量最少。进一步分析表明,该释放量既能保证空心莲子草叶甲的稳定繁育和数量增加,又可获得对空心莲子草最佳的取食和控制效果,具有成本低、效率高的特点。试验结果可为进一步优化空心莲子草叶甲的繁育和释放技术提供理论依据。     

入侵植物空心莲子草的入侵机制及综合管理

综合分析空心莲子草(Alternanthera philoxeroides)的入侵机制并探讨其防治方法,指出空心莲子草的成功入侵与其自身的生物学特性,如无性繁殖、较强的表观可塑性、广泛的适应性、强大的抗逆性等密切相关,其次与生境的可入侵性、人为干扰等外在因素也息息相关。应将人工防除与化学、生物防除手段相结合,加强区域间管理,适时适地采取最佳防除措施。需加强入侵种空心莲子草与本地种的竞争关系研究,运用本地种防止其入侵。     

贵州草海湿地空心莲子草入侵迹地植物群落结构数量特征

草海湿地是中国西南地区极为重要的喀斯特高原高寒湿地生态系统,也是中国黑颈鹤(Grus nigricollis)等珍稀水鸟重要的越冬场所之一。空心莲子草(Alternanthera philoxeroides)已在草海环湖消涨区、航道、码头等地形成入侵趋势,造成湿地植物群落物种多样性降低、破坏湿地生态系统稳定,严重威胁草海湿地生态安全。本研究选择贵州草海湿地空心莲子草入侵迹地植物群落作为研究对象,采用标准样地调查法分别在草海湿地上游、中游、下游设置轻度、中度、重度调查样点,对空心莲子草入侵迹地植物群落组成种类、种群密度、重要值等数量特征进行分析,提出空心莲子草清除后湿地植物群落培育方向,为草海湿地入侵植物迹地修复提供参考。研究结果表明,(1)草海湿地空心莲子草入侵迹地植物群落包括9个科,15种植物,其中禾本科(Gramineae)植物4种,莎草科(Cyperaceae)植物3种,蓼科(Polygonaceae)植物2种,菊科(Asteraceae)、苋科(Amaranthaceae)、香蒲(Typhaceae)、睡菜科(Menyanthaceae)、唇形科(Labiatae)、木贼科(Equisetaceae)植物各1种。(2)草海湿地上游入侵迹地植物群落总密度随入侵程度增加呈升高趋势,而草海湿地中游和下游入侵迹地植物群落密度随入侵程度增加呈降低趋势。(3)空心莲子草入侵迹地次优种、伴生种均以禾本科的李氏禾(Leersi ahexandra)、茭白(Zizania latifolia),莎草科的水葱(Scirpus validus)、荆三棱(Scirpus fluviatilis)为主。空心莲子草清除后的植物群落培育应以李氏禾、水葱、荆三棱、茭白为主要建群种。     

Clonal extent,apical dominance and networking features in the phalanx angiosperm <Emphasis Type="Italic">Zostera noltii</Emphasis> Hornem.

Disaggregating seagrass meadows and studying its components separately (clones, ramets, shoots) can provide us insights on meadow dynamics and growth patterns. The clonal growth, dependent upon clonal rules may regulate and impose constraints to plant architecture and, therefore, determine how individual clones evolve into the environment. In order to investigate the relationship between clonal growth rules and clone architecture, the belowground network architecture of single-clones of the seagrass Zostera noltii was studied. Networks were traced in situ after washing out the overlying sediment, and network characteristics were measured using digital analysis: area covered by clone, total rhizome length, type of rhizomatic axes (main, secondary, tertiary, quaternary), number and length of the internodes, branching angles and branching frequencies. This approach revealed that Z. noltii is able to develop into large clones integrating up to 300 internodes, 676 cm of rhizome, 208 shoots and 4,300 cm² of plant area. Internodal length depended on both, the distance to the apical shoot (time effect) and the axes type (apical dominance effect). However, average branching angle was independent of axis type (average 58.3 ± 0.75), but varied significantly depending on the distance from the apical shoot. This average branching angle allows Z. noltii maximize the rate of centrifugal expansion, maintaining a high density in colonized areas to produce close stands but also minimizing the investment in belowground biomass and ramets overlapping. The clonal architecture of Z. noltii seems to be regulated by the interaction of both, apical dominance strength and clonal integration distance. Moreover, clonal growth rules and growth pattern seem to constrain clonality through (clonal) plant architecture regulations (i.e. branching is restricted in secondary axes, similar average branching angles regardless the axes, the higher the distance to the apex the higher the number of internodes in secondary axes, shorter internodes in secondary and tertiary axes). Future research efforts should focus on how these complex relationships between apical dominance and clonal integration interact to elucidate the temporal (seasonal) and spatial scales of both processes and the outcome at the plant architectural level.     

异质环境下入侵植物薇甘菊的适应性与繁殖特性

薇甘菊（Mikaniamicrantha H.B.Kunth）为世界上最具有入侵性和危害性的外来入侵物种之一,对其控制与管理已成为长期以来世界性难题。了解入侵植物薇甘菊在异质环境下的适应性与繁殖特性对安全有效预警、监测和防治该入侵物种具有重要意义。本文于薇甘菊盛花期,根据薇甘菊入侵生境的光照条件、群落结构和生长方式差异,在薇甘菊常见的4种入侵生境（林地、荒地、农田和和河边）选取10个样地,调查研究了不同生境条件下薇甘菊的开花结实和繁殖分配,并运用植物叶片功能性状的研究方法研究了不同生境条件下薇甘菊的比叶面积（SLA）、叶干物质比例（LMF）和茎干物质比例（SMF）的3种叶片功能性状。研究结果表明,生境条件对薇甘菊的开花结实和繁殖分配具有显著影响。其中抛荒农田（样地Ⅷ）和河流边缘（样地Ⅸ和Ⅹ）薇甘菊的总花数、种子量、生殖枝茎生物量分配和花生物量分配均显著大于其他生境,显示在有利于其生长的条件下,薇甘菊种群倾向于有性繁殖;而在不利于薇甘菊生长的低光照林地（样地Ⅰ）和种间竞争强的农田红薯地（样地Ⅵ）生境条件下,薇甘菊的总花数和种子量明显小于其他生境,但营养枝叶生物量分配显著大于其他样地,而茎生物量分配值则处于中间值,表明在不利于其生长的生境条件下,薇甘菊通过提高营养枝茎的生物量分配和叶的生物量分配来适应,其种群则更倾向于克隆繁殖。通过分析不同生境条件下薇甘菊的叶片功能性状,结果表明,在不利于薇甘菊生长的低光照林地（样地Ⅰ）和种间竞争强的农田红薯地（样地Ⅵ）生境条件下,薇甘菊的叶面积和茎干物质比例（SMF）显著小于其他样地,且彼此差异不显著;但比叶面积（SLA）和叶干物质比例（LMF）则显著大于其他样地,显示薇甘?     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Effects of clonal integration on the growth and physiology of Phalaris arundinacea under simulated herbivory北大核心CSCD

In order to investigate the effects of clonal integration on the adaptation of clonal plants to the environmental stress of homogeneous herbivory, we conducted a greenhouse experiment to investigate the ecophysiological response characteristics of Phalaris arundinacea collected from the riparian zone of the Poyang Lake wetland and Le'an River in Jiangxi Province and the effect of clonal integration on this dominant plant under two-months of simulated cyclical homogenous herbivory. Simulated herbivory strength was set at four different levels: no leaf removal (control) and 25%, 50%, and 75% leaf removal. We implemented two methods of clonal integration including no integration, in which the rhizome connection was severed, and integration, in which the rhizome connection was intact. We found that simulated herbivory significantly decreased the number of leaves, total shoot length, number of ramets, and biomass of P. arundinacea (P < 5%), regardless of whether the rhizomes were intact or severed. Ramets with severed rhizome connections (RRC) generally had higher growth indices than those with intact rhizome connections (RIC). Severing rhizomes did not significantly affect the chlorophyll content of P. arundinacea, while RRC under simulated herbivory intensities of 25% and 50% leaf removal had higher chlorophyll a (Chl a), chlorophyll b (Chl b), and total chlorophyll (Chlt) contents than RIC did, and simulated herbivory increased the chlorophyll contents of all ramets. RRC net photosynthesis rate (Pn), stomatal conductance (Gs), intercellular CO2 concentration (Ci), and transpiration rate (Tr) were all higher in RRC than they were in RIC under simulated herbivory intensities of 50% and 75% leaf removal; however, the opposite relationship was observed under a simulated herbivory intensity of 25% leaf removal. Further, simulated herbivory limited the photosynthetic index of P. arundinacea. These effects resulted in an inadequate accumulation of nutrients in the plant. Phalaris arundinacea can adapt to simulated herbivory treatments in terms of growth, but clonal integration cannot improve the growth of P. arundinacea under a homogeneous herbivory treatment. © 2018 Science Press. All rights reserved.     

Demographic feedback between clonal growth and fragmentation in an invasive seaweed

Many abundant plants, invertebrates, and seaweed are clonal, and this allows the formation of high-density aggregations, foraging, and the placement of modules into new space, and rapid rates of expansion. For these species, population density and rates of expansion are functions of recruitment of asexual modules and post-recruitment vegetative growth and survivorship. In this study, we provide the first experimental test of the relative importance of these two processes in determining the abundance of a clonal seaweed using Caulerpa taxifolia, an invasive green alga that spreads rapidly and reaches very high abundance. We asked two main questions: What is the relative importance to abundance (biomass) of vegetative stolon growth and fragment recruitment during expansion of established patches? Does greater fragment recruitment result in greater abundance in established patches? Vegetative growth of stolons underpinned patch expansion. Plots with stolons growing into them always had a greater abundance than plots where stolons were removed, even when fragment recruitment was increased. Greater recruitment only resulted in greater abundance when stolons were absent, a situation analogous to the establishment of new populations. Although post-recruitment processes were more important in determining abundance during patch expansion, there was greater ambient fragment recruitment when stolons were present compared to when they were absent, and as the abundance of C. taxifolia increased, demonstrating an important feedback between stolon growth, abundance, and fragment recruitment. In established patches, greater fragment recruitment over six months (six levels ranging from 0 to 480 recruits x m(-2) x mo(-1)) had no effect on biomass. Our experiments demonstrate that the rapid expansion and high abundance of invasive C. taxifolia are underpinned by post-recruitment vegetative growth and, during expansion, by a feedback between vegetative growth and asexual fragmentation.     

林下和撂荒地生境紫茎泽兰幼苗的生长特征及其种内竞争形态反应

比较研究了林下和撂荒地生境有无竞争条件下紫茎泽兰（Ageratina adenophora（Sprengel））幼苗的生长特征、生长速率、生物量和种内竞争的形态反应。结果表明：（1）2类生境条件下紫茎泽兰植株幼苗形态特征基本一致,无竞争条件下紫茎泽兰幼苗根长和叶片数明显高于在竞争条件下对应指标值;（2）同类生境中,在无竞争条件下紫茎泽兰幼苗的生长速率、生物量积累显著比在竞争条件下的高;（3）不同生境中,撂荒地生境紫茎泽兰幼苗生物量显著高于林下生境的值,在无竞争条件下,林下生境紫茎泽兰幼苗生长速率比撂荒地生境的高,而在竞争条件下反之;（4）林下和撂荒地生境紫茎泽兰幼苗植株竞争强度差异不显著,但在林下生境其地上部分的竞争强度显著大于其在撂荒地生境的值,而根的竞争强度相反。可见,紫茎泽兰幼苗在种群建立的过程中具有不同的生态适应性和形态反应,这可能是其能成功入侵不同生境的一种重要生理生态策略。     

三叶鬼针草生物量分配与化感作用对大气温度升高的响应

全球气候变暖已经成为不容置疑的事实,同时外来入侵植物对入侵地的生态环境造成严重的危害,外来入侵植物可能对温度升高做出积极地响应。文章研究了不同温度（22、26和30℃）处理对入侵植物三叶鬼针草（Bidens pilosa L.）种子萌发、幼苗生物量分配及化感作用的影响,探讨三叶鬼针草对全球气候变暖的响应策略。结果表明;温度为22和26℃比30℃有利于三叶鬼针草种子的萌发。温度升高显著增加三叶鬼针草的株高、生物量和叶面积,三叶鬼针草幼苗增加对茎和叶的生物量投资。同时相同浓度的三叶鬼针草水浸提液对马唐（Digitaria sanguinalis（L.）Scop）和牛筋草（Eleusine indica（L.）Gaertn）的化感作用随着温度升高而增强。研究表明：温度升高促进了三叶鬼针草的生长,改变生物量分配模式同时增强了对受体植物的化感作用。温度升高可能是促进三叶鬼针草入侵的生态环境因子之一,未来全球气温变暖可能使其入侵加剧。     

Optimal biomass allocation in heterogeneous environments in a clonal plant—Spatial division of labor

When interconnected ramets of clonal plants are growing in heterogeneous environments, ramets may specialize to uptake locally abundant resources rather than scarce resources. This biomass allocation pattern may result in more efficient sharing of resources through physiological integration and an overall benefit to the plants (spatial division of labor; DoL).