Is the removal of aboveground shrub biomass an effective technique to restore a shrub‐encroached grassland?

Encroachment of woody plants into grasslands is a global phenomenon that has substantial impacts on pastoral productivity and ecosystem services. Over the past half century, pastoralists and land management agencies have explored various options to control woody plants in order to improve ecosystem services in shrub‐encroached grasslands. We examined the effectiveness of controlling the encroachment of the shrub Caragana microphylla into grassland in Inner Mongolia, China. We cut and removed all of the aboveground biomass from 450 shrubs, predicting that the effectiveness of this technique to control shrubs would depend on shrub morphology. Specifically, we expected that larger shrubs with more biomass would be more difficult to kill by cutting than smaller shrubs. A year after treatment, we found that cutting killed only 11% of the 450 treated shrubs, and of these, three‐quarters of the locations that they occupied reverted to grasses and one‐quarter to bare soil. Shrubs that survived the cutting treatment produced more stems and leaf biomass, and therefore had a greater leaf to stem ratio. Shrubs that died after cutting had a lower crown area and basal area, and less stem biomass than shrubs that resprouted within 12 months of cutting. There were no effects of shrub height on the fate of treated shrubs. Cutting had no effect on understory plant cover or richness, but reproductive plants were taller under shrubs that were not cut. Overall, our study showed that removing aboveground shrub biomass by cutting is an ineffective technique for “restoring” the original grassland community unless shrubs are very small. Strategic targeting of small shrubs would be a more effective technique for controlling the spread of C. microphylla in the long term.     

Effect of springtime solar ultraviolet-B radiation on growth of Colobanthus quitensis at Palmer Station, Antarctica

We examined the influence of solar ultraviolet‐B radiation (UV‐B; 280–315 nm) on the growth of Colobanthus quitensis plants by placing them under contrasting UV‐B filters at Palmer Station, along the Antarctic Peninsula. The filters reduced diurnal biologically effective UV‐B (UV‐B_BE) either by 83% (‘reduced UV‐B’) or by 12% (‘near‐ambient UV‐B’) over the 63 day experiment (7 November 1998–8 January 1999). Ozone column depletion averaged 17% during the experiment. Relative growth and net assimilation rates of plants exposed to near‐ambient UV‐B were 30 and 20% lower, respectively, than those of plants exposed to reduced UV‐B. The former plants produced 29% less total biomass, as a result of containing 54% less aboveground biomass. These reductions in aboveground biomass were mainly the result of a 45% reduction in shoot biomass, and a 31% reduction in reproductive biomass. Reductions in shoot biomass were owing to an 18% reduction in branch production by main shoots, while reductions in reproductive biomass were the result of a 19% reduction in individual capsule mass. Total plant leaf area was reduced by 19% under near‐ambient UV‐B, although total leaf biomass was unaffected because leaves had a greater specific leaf mass. The reduction in plant leaf area under near‐ambient UV‐B was attributable to: (1) production of 11% fewer leaves per main shoot system and plant, which resulted from an 18% reduction in branch production by main shoots. Leaf production per individual main shoot or branch was not affected; (2) shorter leaf longevity—main shoots contained 14% fewer green leaves at a given time; and (3) smaller individual leaves—leaf elongation rates were 14% slower and mature leaves were 13% shorter.     

Stem hydraulic properties of vines vs. shrubs of western poison oak,Toxicodendron diversilobum

Summary This study investigated the effect of mechanical support on water transport properties and wood anatomy of stems of western poison oak, Toxicodendron diversilobum (T. & G.) Greene. This plant grows as a vine when support is present but as a shrub when support is absent. I compared vines and shrubs growing naturally in the field and those produced from cuttings of 11 source plants in a common garden. Huber value (xylem transverse area/distal leaf area) was lower but specific conductivity (water volume · time^-1 · xylem transverse area^-1 · pressure gradient^-1) was higher in supported than unsupported plants both in the field and the common garden. The opposing effects of Huber value and mon garden. The opposing effects of Huber value and specific conductivity resulted in the same values of leafspecific conductivity (LSC, water volume · time^-1 · distal leaf area^-1 · pressure gradient^-1) for supported and unsupported shoots at a given site. Therefore, for the same rates of evapotranspiration, supported and unsupported shoots will have the same pressure gradients in their stems. Vessel lumen composed a higher proportion of stem cross-section in supported than unsupported plants (due to slightly wider vessels and not to greater vessel density). These results suggest that the narrow stems of supported plants are compensated hydraulically by the production of wider vessels: at a given site, poison oak plants co-ordinate their leaf and xylem development such that their stems achieve the same overall conductive efficiencies (LSCs), regardless of support conditions.     

Phenology and resource allocation in a high arctic evergreen dwarf shrub, Cassiope tetragona

During the growing season of 1981, we studied phenological patterns of development of vegetative shoots and sexual organs, partitioning of biomass and productivity, and partitioning and concentrations of N, P, K, Ca, and Mg for the evergreen dwarf shrub Cassiope tetragona (Ericaceae) at a high arctic lowland oasis at 79°N on Ellesmere Island, Canada. Short elongation (3.6 mm yr⁻¹) and net production (5% of aboveground phytomass) of Cassiope were small. Vegetative and reproductive development were initialed just after snowmelt. Seasonal patterns of allocation of phytomass and nutrients to current growth were characterized by gradual accumulation of quantity over the growing season. However, concentrations of N, P, and K decreased by ca 50% in current tissues during the growing season, while Ca increased with tissue age. Overwinter storage of nutrients occurred in live aboveground shoots. The large proportion of phytomass that occurred as aboveground, attached dead material, as well as the small net production and belowground standing crop, suggest an allocation pattern more similar to that of high arctic cushion plants, than to other evergreen, ericaceous dwarf shrubs occurring at lower latitudes in the Arctic.     

C3 shrub expansion in a C4 grassland: positive post-fire responses in resources and shoot growth

Changes in land management and reductions in fire frequency have enabled woody species to increase in grasslands worldwide. Nevertheless, fire is rarely eliminated from grasslands, and for shrubs to survive, they must be able cope with fire and replace aboveground structures. Because new shoots may have more available solar radiation, greater root?:?shoot ratios, and thus more resources available belowground after fire compared to undisturbed shrub communities, we hypothesized that carbon, nutrient, and water relations may be enhanced in stems compared to those in an undisturbed grassland. However, this same post-fire resource pulse stimulates the grasses and may intensify competitive interactions between shrubs and grasses. To test these predictions, we measured seasonal patterns in net photosynthesis (A), predawn xylem pressure potentials (XPP), leaf nitrogen (N) content, and productivity of Cornus drummondii shoots from shrub patches (islands) of different sizes in mesic grasslands burned annually, burned infrequently, and protected from fire. Seasonal average A was 20% higher (P = 0.016) in burned than in unburned shrubs, regardless of island size. Shrubs in burned sites also produced shoots with higher leaf N than unburned shrubs, and N content was higher in leaves from small islands compared to large islands (P < 0.0001). Burning caused a decrease in late summer predawn XPP in small islands (-3.1 MPa), whereas burned large islands did not differ from unburned shrubs. Post-fire productivity of new shoots was significantly greater compared to shoots in unburned sites. These results indicate that a transient period of high resource availability after fire allows for increased growth and rapid recovery of grassland shrubs. Thus, although fire has a negative effect on aboveground biomass of shrubs, the post-fire increases in resource availability, which enhance growth in the dominant grasses, are also important for recovery of woody species.     

Biomass estimation models and allocation patterns of 14 shrub species in Mountain Luya,Shanxi, China

Aims Shrub species have evolved specific strategies to regulate biomass allocation among various organs or between above- and belowground biomass and shrub biomass model is an important approach to estimate biomass allocation among different shrub species. This study was designed to establish the optimal estimation models for each organ (leaf, stem, and root), aboveground and total biomass of 14 common shrub species in Mountain Luya, Shanxi Province, China. Furthermore, we explored biomass allocation characteristics of these shrub species by using the index of leaf biomass fraction (leaf to total biomass), stem biomass fraction (stem to total biomass), root biomass fraction (root to total biomass), and root to shoot mass ratio (R/S) (belowground to aboveground biomass).
Methods We used plant height, basal diameter, canopy diameter and their combination as variables to establish the optimal biomass estimation models for each shrub species. In addition, we used the ratios of leaf, stem, root to total biomass, and belowground to aboveground biomass to explore the difference of biomass allocation patterns of 14 shrub species.
Important findings Most of biomass estimation models could be well expressed by the exponential and linear functions. Biomass for shorter shrub species with more stems could be better estimated by canopy area; biomass for taller shrub species with less stems could be better estimated by the sum of the square of total base diameter multiply stem height; and biomass for the rest shrub species could be better estimated by canopy volume. The averaged value for these shrub species was 0.61, 0.17, 0.48, and 0.35 for R/S, leaf biomass fraction, stem biomass fraction, and root biomass fraction, respectively. Except for leaf biomass fraction, R/S, stem biomass fraction, and root biomass fraction for shrubs with thorn was significantly greater than that for shrubs without thorn.     

Nutrient and carbon relations in subalpine dwarf shrubs after neighbour removal or fertilization in northern Italy

Two subalpine dwarf-shrub heath communities with differing levels of soil nutrient availability were subjected to a 3-year experimental manipulation, including nutrient addition or removal of one of the two co-dominant species from each community. The main objective of our study was to assess the relative importance of interspecific competition versus nutrient limitation in relation to soil fertility. We also aimed to investigate if and to what extent current-year shoot size, leaf-based rates of net photosynthesis and foliar nutrient status accounted for the observed changes in the aboveground biomass of the shrubs. At the end of the experiment, neighbour removal increased the aboveground biomass of all shrubs, especially in the more fertile community, while fertilization did not. We concluded that: (1) competition is more effective than nutrient limitation in structuring the vegetation of subalpine heathlands; and (2) competition intensity is stronger in the more fertile community. The observed patterns of variations in aboveground biomass were not consistently related to net photosynthetic rates, size of individual shoots and foliar nutrient status. Hence, we also concluded that the growth response of dwarf shrubs to altered environmental conditions is primarily determined by developmental plasticity.     

Photosynthetic and biomass allocation responses of Liquidambar styraciflua (Hamamelidaceae) to vine competition

This 2-year field study examined stomatal conductance, photosynthesis, and biomass allocation of Liquidambar styraciflua saplings in response to below- and aboveground competition with the vines Lonicera japonica and Parthenocissus quinquefolia. Vine competition did not affect stomatal conductance of the host trees. The leaf photosynthetic capacity and photosynthetic nitrogen-use efficiency were significantly reduced by root competition with vines, either singly or in combination with aboveground competition, early in the second growing season. However, such differences disappeared by the end of the second growing season. Trees competing below ground with vines also had lower allocation to leaves compared with steins. Aboveground competition with vines resulted in reduced photosynthetic capacity per unit leaf area, but not per unit leaf weight, in trees. No correlation was found between single leaf photosynthetic capacity and tree growth. In contrast, a high positive correlation existed between allocation to leaves and diameter growth. Results from this study suggest that allocation patterns are more affected than leaf photosynthesis in trees competing with vines.     

东北山地灌木沼泽植物铜、锌分布特征及季节动态

研究了东北山地灌木沼泽主要植物--细叶沼柳、五蕊柳、丛苔草和修氏苔草中Cu、Zn分布、积累及其季节动态.结果表明:Cu含量变动范围为6～12 mg·kg-1,细叶沼柳和五蕊柳各器官Cu含量为根》枝》叶;丛苔草和修氏苔草为茎》叶》根,Cu主要积累在灌木的根系和苔草的茎叶中,灌木和苔草中Cu含量相差较小.Zn含量变动在30～250 mg·kg-1之间,细叶沼柳和五蕊柳各器官Zn含量为叶》枝》根,尤其是叶和枝中都在150 mg·kg-1以上;丛苔草和修氏苔草各器官Zn含量为根》茎》叶.Zn多积累在灌木的叶和苔草的根中,且灌木各器官Zn含量明显高于苔草.细叶沼柳和五蕊柳各器官对Zn的富集系数均大于1.45,显示出较强的Zn富集能力.4种供试植物在生长初期地上部分Cu、Zn含量较高,且随着季节变化呈波动式降低趋势,而根中则表现出生长初期和末期Cu、Zn含量较高的特点.     

Does morphological plasticity of the <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> canopy increase invasiveness?

Morphological plasticity could facilitate invasions of wetland plants into areas that experience increased durations of flooding and eutrophication. We explored canopy plasticity of Phalaris arundinacea, an aggressive invader of wetlands, as it differentially invaded wet prairie mesocosms under 3 flooding durations and 3 levels of nutrient addition. Phalaris grew as a sward with intermittent and early-season flooding but shifted to tussocks under constant flooding. These two growth forms differed by >20% in several canopy ratios. Clones that formed tussocks produced 45% more shoots per unit biomass (P = 0.007) and a 25% higher ratio of total shoot length to biomass (P = 0.04). Lighter-weight shoots supported 33% fewer leaves and, consequently, had 35% less leaf area per shoot height (P < 0.002). Tussocks developed a continuous mat of adventitious roots, with root mats reaching 20.9 ± 0.6 cm in diameter and 4.7 ± 0.3 cm in height over two growing seasons. While forming tussocks, Phalaris tolerated longer durations of flooding and more than doubled its aboveground biomass. Invasions occurred rapidly, with Phalaris exceeding 75% canopy cover and accounting for 66% of the total aboveground biomass under constant flooding. Early-season flooding increased the lateral spread of individual shoots. High nutrient addition produced shoots that were 27% taller and 50% heavier (P < 0.02), with 81% more leaf area (P < 0.0003) than shoots that received no nutrients. Consequently, under early-season flooding with high nutrient additions, Phalaris was primed to invade, nearly doubling its proportion of the total aboveground biomass and exceeding 50% canopy cover during year two.     

不同生境和森林内薇甘菊的生存与危害状况

对薇甘菊（Ｍｉｋａｎｉａ ｍｉｃｒａｎｔｈａ Ｈ．Ｂ．Ｋ）在不同生境下生长状况及其对其它植物的危害程度的研究表明,这种热带杂草是喜光、好湿的。薇甘菊伴生物种以藤本植物居多。其中葛藤和五爪金龙出现的频度最大。薇甘菊及其伴生种通过攀爬树冠,形成盖幕作用而对其它植物造成危害。在森林内薇甘菊危害的对象主要是低矮的乔木对高大乔木影响不大。     

Why vines have narrow stems: Histological trends in Bauhinia (Fabaceae)

Summary Xylem (wood) tissue in plants functions both for mechanical support and water transport. Since vines are mechanical parasites, they allocate less biomass for xylem tissue than do free-standing trees or shrubs. With-in the genus Bauhinia, stems of vine species were found to have not only less xylem per distal leaf area, but also less phloem and cortical tissue when compared to tree and shrub species. The phloem and cortical reductions are interpreted as an indirect effect of the developmental/geometric constraints imposed by the evolution of a reduced mechanical system. Apparently vines overcame these constraints with the evolution of wider vessels and wider sieve tubes and with many types of variant (anomalous) secondary growth. The long and wide vessels of vines, which compensate hydraulically for the reduced xylem areas, may help limit the distribution of vine species in nature.     

放牧对内蒙古锡林河流域草地群落植物茎叶生物量资源分配的影响

以内蒙古锡林河流域沿水分梯度分布的灰脉苔草(Carex appendiculata)、贝加尔针茅(Stipa baicalensis)、羊草(Leymus chinensis)、大针茅(Stipa grandis)、小叶锦鸡儿(Caragana microphylla)和冷蒿(Artemisia frigida) 6个草地群落为对象, 研究了围封禁牧与放牧样地中144个共有植物种的高度、丛幅面积、茎、叶和株(丛)生物量、茎叶比等性状。结果表明: 1)在个体水平上, 放牧样地中植物的生殖枝高度、营养枝高度、丛幅面积、单株(丛)生物量、茎、叶生物量和茎叶比均显著低于围封禁牧样地, 植物在放牧干扰下表现出明显的小型化现象; 2)在群落水平上, 放牧亦显著降低了群落总生物量和茎、叶生物量; 3)过度放牧显著改变了物种的资源分配策略, 使生物量向叶的分配比例增加, 向茎的分配比例减少。资源优先向同化器官分配可能是植物对长期放牧干扰的一种重要适应对策; 4)轻度放牧对物种的资源分配没有显著影响, 单株(丛)生物量和群落茎、叶及总生物量均表现出增加趋势, 这与过度放牧的影响正好相反。过度放牧引起的植物个体小型化改变了生态系统中物种的资源分配策略, 进而对生态系统功能产生重要的影响。     

扰动环境中不同刈割方式柠条营养生长补偿的影响

 柠条（Caragana korshinskii）在地上组织破坏后进行补偿性生长,这是重复利用柠条资源的基础,但对 柠条不同刈割方式下营养生长补偿的模式有待探讨。该文通过5种刈割方式：去除主枝长的30%（30%RSL） 、去除主枝长的60%（60%RSL）、去除分枝数的25%（25%RSN）、去除分枝数的50%（50%RSN）和去除分枝 数的100%（100%RSN）来研究柠条的营养生长补偿。结果表明：刈割处理的柠条生物量当年发生了超补偿 ,当年生枝数/枝、当年生枝长、当年生枝生物量/枝、当年生枝生物量/株比对照高。对照、30%RSL和 60%RSL 处理未长出基梢。 100%RSN处理的基梢数/刈割枝、基梢长、单个基梢平均生物量显著高于25%RSN 和50%RSN处理,基梢生物量/株随刈割去除生物量的增加而增加。100%RSN 处理未结果,其它处理果实产量 表现出超补偿或精确补偿。对照处理营养生长和生殖生长均低,其它处理当年生枝生物量与果实产量成显 著负相关。从整个生长季节来看,营养生长主要集中在果实成熟之前。我们认为,100%RSN处理是柠条地上 组织破坏后尽快恢复的合理方式,其当年生生物量远高于其它处理。顶端优势的破坏促使休眠芽的萌发, 根冠比的改变使地上组织获得较多养分和水分,根系储存的碳水化合物的供应是促使刈割柠条营养生长超 补偿的的可能机制,而减少生殖生长对资源的消耗,是100%RSN处理地上生物量尽快恢复的另一重要因素 。     

SEXUAL DIMORPHISM IN BIOMASS ALLOCATION AND CLONAL GROWTH OF XANTHOXYLUM AMERICANUM

Reproduction can have a high resource cost. It has been suggested that greater investments in sexual reproduction by female dioecious plants leads to a lower rate of vegetative growth in females than in males. In this study, we investigated sexual dimorphism in biomass allocation and genet growth of the dioecious clonal shrub, northern prickly ash (Xanthoxylum americanum). The allocation of biomass over the course of one growing season to reproductive tissue, leaves, and growth of aboveground first-year wood, was compared in 18 clones growing in fields and six clones in woods in southeastern Wisconsin during 1985 and 1986. In addition, the number of shoots per clone, and weight of nonfirst-year wood (accumulated biomass) above- and below-ground were estimated. In open field sites, male clones allocated more biomass to new wood and less to reproduction than females, although males allocated more to flowers alone. Accordingly, male clones had significantly more shoots and more accumulated biomass both above- and below-ground than female clones. In the woods, where fruit set was near zero, there were few significant differences between male and female clones in either biomass allocation or accumulated biomass. These results support the hypothesis that the high resource investment in fruit production by females reduces their vegetative growth relative to males.     

江西常绿阔叶林木本植物不同冠层高度当年生小枝茎构型对叶生物量的影响

以江西阳际峰自然保区69种木本植物为对象,采用标准化主轴回归(SMA)的方法,对不同冠层高度当年生小枝的构件生物量(叶生物量、茎生物量、小枝生物量)与茎构型特征(茎直径、茎长度、茎宽长比、茎体积和茎密度)进行了分析,研究当年生小枝茎构型对叶生物量的影响.结果表明:不同冠层高度及生活型间叶生物量、茎生物量、小枝生物量、茎直径、茎长度、茎宽长比及茎体积均无显著差异,而茎密度差异显著.不同冠层高度及生活型间,叶生物量与茎生物量及总生物量之间均呈显著等速生长关系.小枝叶生物量分别与茎直径、茎体积呈显著异速生长关系,且该异速生长指数在不同冠层高度无显著差异.茎长度、茎宽长比及茎密度对当年生小枝叶生物量变异的解释力较小(<24%).冠层高度和生活型对小枝叶-茎生物量的等速分配关系影响不显著.小枝茎构型中,相比于茎长度、茎宽长比以及茎密度,茎的直径与体积对当年生小枝叶生物量的影响更大,冠层高度对当年生小枝茎构型与叶生物量间的异速分配模式无显著影响.     

Comparisons among biomass allocation and spatial distribution patterns of some vine,pteridophyte, and gymnosperm shoots

The interspecific allometry of leaf, stem, and reproductive biomass distal to stem diameter was determined for a total of 12 angiosperm vine, gymnosperm, and pteridophyte species to compare allocation patterns to vegetative and reproductive shoot organs. The allometry of stem diameter in terms of the distance from shoot apices also was determined to quantify the manner in which vines, gymnosperms, and pteridophyte stems tapered along their length. The stems of vine species were found to weigh more than those of arborescent gymnosperm species distal to any point of equivalent stem diameter. Vine species also distribute more of their stem mass to shoot length as opposed to girth than gymnosperm species. Vine stems also supported proportionally larger leaf and reproductive biomass in comparison to gymnosperm stems of equivalent diameter, yet partitioned their total shoot biomass more or less equally between leaf and stem biomass in the same manner as the gymnosperm species examined. The allometry of vine as well as gymnosperm leaf biomass with respect to stem biomass appeared to be slightly anisometric and negative, suggesting that more massive stems had proportionally less leaf biomass than their smaller, less massive counterparts. Vine stems could be approximated as very slender cones; the shape and geometry of gymnosperm stems complied with those of stubby, truncated cones whose top diameter (for those examined), on the average, equaled 28% of the basal diameter. In general terms, the interspecific allometry of vines was most similar to that of pteridophytes. Collectively, these data refute the commonly held notion that vine stems are simply more slender than those of species with self-supporting stems.     

Effects of imazamethabenz on the main shoot growth and tillering of wild oat (Avena fatua L.)

Foliar application of imazamethabenz at sublethal doses of 100 and 200 g a.i./ha to wild oat plants at the two-leaf stage without tillers greatly inhibited the growth of the main shoot but increased tillering. The near cessation of sheath and the main stem elongation indicated that the major effect of imazamethabenz on the main shoot was inhibition of intercalary growth. Low doses of imazameth-abenz treatment resulted in more leaves (including leaf primordia) in the main stem but did not affect mature first and second leaves. Sublethal doses of imazamethabenz only briefly inhibited tiller growth. A later increase in tillering in treated plants resulted from the stimulated resumed growth of tillers and the increased initiation of tiller buds. Such enhanced tillering mainly resulted from the release of apical dominance due to the inhibition or cessation of the main stem growth with imazamethabenz treatment. Both doses of imazamethabenz (100 and 200 g a.i./ha) significantly reduced the biomass of shoots and roots, but increased the ratio of roots/ shoots dry weight.     

Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.