Restoration Ecology of an Endangered Plant Species: Establishment of New Populations of Cirsium pitcheri

We determined the effects of shade, burial by sand, simulated herbivory, and fertilizers on the survival and growth of artificially planted population of Cirsium pitcheri—an endangered plant species of the sand dunes along Lake Huron. Sand burial experiments showed that greenhouse grown plants should optimally be transplanted into areas receiving 5 cm of sand deposition: burial at this depth maximized emergence, survivorship, and below‐ground biomass. Under field conditions, simulated herbivory of up to 50% of the plant height produced a slight increase in biomass after one year of growth. Field observations showed that when white‐tailed deer removed more than 50% of the transplant's leaf tissue, the plant died. The application of a 20:20:20 (N:P:K) water‐soluble fertilizer produced a significant increase in the dry leaf biomass, total leaf area, and total dry biomass relative to control plants. We also tested for the presence or absence of a persistent seed bank. Few seeds were recovered from soil samples collected from Pinery Provincial Park and Providence Bay. However, C. pitcheri has the ability to form a persistent seed bank under field conditions but only at soil depths of 15 cm. Cirsium pitcheri seeds are able to germinate and seedlings can emerge from a burial depth of up to 6 cm. Thus, seeds planted in open, sunny areas will probably maximize emergence, growth, and survivorship of seedlings. Populations of C. pitcheri can be restored by planting seeds at shallow depths, transplanting greenhouse‐grown plants, applying water soluble fertilizers, and protecting plants from herbivores.     

Seedling growth variation in response to sand burial in four <Emphasis Type="Italic">Artemisia</Emphasis> species from different habitats in the semi-arid dune field

We compared seedling growth of four Artemisia species dominated at different habitats to determine whether interspecific seedling growth variation of a same genus in tolerance to burial can be used to explain plant distribution in the sand dune field. Interdune lowland species, Artemisia gmelinii, stabilized dune species, A. frigida, semi-stabilized dune species, A. halodendron, and active dune species, A. wudanica were selected. Seedlings grown for 3 weeks were treated at five burial depths for three burial times in pot experiments. Species from the habitats with little burial had smaller survival rate, dry weight and stem elongation speed than those from the habitats with intensive burial when buried. Furthermore, when buried, the former tended to adjust biomass allocation between shoot and root and produce adventitious buds, while the latter tended to maintain a constant root:shoot ratio and produce adventitious roots. We conclude that (1) seedlings of species with a long evolutionary history of exposure to sand burial (from the active sand dune), show quicker stem growth when buried than do seedlings of species from the habitats with little or no sand burial; (2) seedlings of species which do not change root:shoot ratio might be more tolerant of sand burial than those do; (3) seedlings of species from the habitats with intensive sand burial is prone to produce adventitious roots and seedlings of species from the habitats with little or no sand burial tend to produce adventitious buds when buried.     

Responses of Hedysarum Laeve , a guerrilla clonal semi-shrub in the Mu Us sandland, to local sand burial

In arid and semi-arid inland deserts, one of the environmental stresses for plants is recurrent sand burial, which can influence the physical and biotic microenvironments of the plants and soil. Previous studies have shown that different levels of sand burial have different effects on plants. Slight sand burial could increase the height increment, leaf biomass and the number of new ramets of the plants while heavy sand burial could impair the growth of the plants and even decrease their chances of survival. In other words, below a certain threshold level of burial, the growth of plants is stimulated probably because of multiple factors. However, as the level of burial increases, the positive response starts to decline until it becomes a negative value. Arid and semi-arid inland deserts are frequently colonized and stabilized by many rhizomatous clonal plants. Clonal physiological integration often helps clonal plants buffer local environmental stress encountered by ramets. A rhizomatous clonal semishrub, Hedysarum laeve (H. laeve), is the dominant plant species and important for vegetation restoration in the Mu Us sandland. To investigate whether clonal integration can increase the threshold of sand burial and help rhizomatous H. laeve tolerate heavy sand burial, we conducted a field experiment. The results showed that slight sand burial could accelerate ramet growth and enhance leaf biomass, stem biomass and shoot biomass, while heavy sand burial reducesed the biomass of the plant and impairs survival and growth of the ramets. Clonal integration increased the threshold of sand burial. Under heavy sand burial, ramets connected to other ramets not buried in sand were more in terms of height increment, stem biomass, leaf biomass and shoot biomass compared to the ramets encountering sand burial but disconnected from other ramets. It suggested that clonal physiological integration could help H. laeve ramets tolerate relatively heavy sand burial. We also discussed that clonal integration plays a role in H. laeve presence in the Mu Us sandland. __________ Translated from Journal of Plant Ecology (formerly Acta Phytoecologica Sinica), 2006, 30(2): 278–285 [译自: 植物生态学报]     

GENETIC ORGANIZATION AND EVOLUTIONARY HISTORY IN TWO NORTH AMERICAN SPECIES OF CIRSIUM

Relationships involving progenitor and derivative species pairs have been documented for a number of plant taxa. In this study, we used starch-gel electrophoresis to examine the probable relationship between Cirsium pitcheri, an endemic thistle restricted to the shorelines of the Great Lakes, and Cirsium canescens, a closely related species of the central Great Plains (U.S.A.). The species shared the same enzyme loci but differed greatly in levels of genetic diversity. C. pitcheri was genetically much less variable than its congener, with fewer polymorphic loci and significantly lower heterozygosity. The genetic variation in C. pitcheri represents a depauperate subset of the alleles present in C. canescens, consistent with the hypothesis that the two species represent a progenitor-derivative species pair. We suggest that C. pitcheri migrated into newly created dune habitats in the Great Lakes region near the end of the Wisconsin glaciation and that its low genetic variability results from repeated and probably prolonged population bottlenecks which accompanied its movement.     

Strong regeneration ability from rhizome fragments in two invasive clonal plants (Solidagocanadensis and S. gigantea)

The two rhizomatous perennials Solidago canadensis and S. gigantea belong to the most widespread alien plants in Europe. Anecdotal observations suggest that they disperse also by rhizome fragments. For testing their resprouting ability, rhizome fragments of different sizes from both species were buried at four different soil depths (0, 5, 10 and 20 cm, respectively) in a common garden. Rhizome fragments of S. canadensis ranged 3–6 cm in length, those of S. gigantea 5–20 cm in length. Resprouting plants were harvested after 3 months and growth related traits measured to assess their vitality. Resprouting rate in S. gigantea was far higher than in S. canadensis (85 and 19%, respectively). In S. gigantea, fragments of all sizes resprouted from all soil depths whereas none rhizome of S. canadensis emerged from 20 cm burial depth. In S. gigantea, size related traits showed significant interactions between fragment size and burial depth, but not relative shoot growth rate. At all burial depths, vitality of plants emerging from small rhizomes was lower than plants emerging from large rhizomes. Effects of rhizome size became stronger with increasing burial depth. The results show that both species are able to resprout from buried rhizome fragments, and that succesful regeneration is more likely to occur in S. gigantea. Managing these species should avoid any activities promoting rhizome fragmentation and dispersal of fragments.     

泥沙掩埋和干旱对三峡库区消落带狗牙根生长恢复的影响

三峡库区消落带植物恢复不仅面临长期淹水逆境,还面临泥沙、干旱等环境因素的胁迫。蓄水期后引种三峡库区消落带狗牙根,掩埋于不同粒径和埋深的沉降泥沙,探讨泥沙掩埋、干旱对反季节淹水后狗牙根出苗和生长恢复的影响。结果表明:在埋深≥8cm和粒径1.25 mm条件下消落带狗牙根萌发苗不能穿透覆盖层出苗。泥沙掩埋导致狗牙根出苗率显著降低,随着埋深增加狗牙根出苗率迅速下降;并且在埋深相同的条件下随着粒径减小狗牙根出苗速率呈增大趋势。随着埋深增加和粒径减小处理组株高、叶长的增长显著被抑制,处理组的株高、叶片数、叶长显著低于对照组。随着干旱、泥沙掩埋复合胁迫时间延长狗牙根分株株高、叶片数、叶长、叶宽的生长发育显著被抑制,且随着泥沙粒径减小和埋深增加其光合电子传递速率呈显著降低趋势,热耗散量显著增大。同时,干旱、泥沙掩埋复合胁迫导致处理组分株光合电子传递速率和热耗散能力显著低于对照组,光合作用能力显著降低,易受到夏季高光伤害。因此,三峡库区周年淹水导致狗牙根对泥沙掩埋以及干旱复合胁迫的耐受能力显著降低,在泥沙掩埋以及干旱复合胁迫下消落带狗牙根的生长恢复能力随粒径减小和埋深增加呈显著降低趋势,会导致库区消落带狗牙根种群衰退演替加快。     

Morphological adaptations byGlehnia littoralis to the biomass and heights of peripheral herbaceous plants in coastal sand dunes

To clarify the effects of peripheral herbal plants onGlehnia littoralis growth in coastal sand dunes, the morphology of their aboveground portions was surveyed in five communities:Carex kobomugi, Calystegia soldanella, Ischaemum anthephoroides, Oenothera biennis, andElymus mollis. Correlation coefficients (CC) were generally significant at the 1% level between community properties [total aboveground biomass (B) and height (H) of dominant species per unit area] and those ofG. littoralis [leaf number (Nl), petiole angle (Anp), petiole length (Lp), petiole weight (Wp), Lp/Wp, Lp/weight of leaf blade (Wb), Wp/total weight (Wt), specific leaf area (SLA), stem length (Ls), and Ls/weight of stem (Ws)J The exceptions were among four pairings: B and NI, B and Wt, H and Nl, and H and Wt. Of the two community properties, biomass had the greatest association with leaf properties while H was most closely related to those of the stems. Petiole angle increased along with leaf order, from 0° to 42° for the C.kobomugi community, from 5° to 55° forCalystegia soldanella, from 49° to 74° forI. anthephoroides, from 54° to 80° forO. biennis, and from 75° to 85° forE. mollis. In all communities, the properties of Wp, SLA, and Wb increased up to the third or fourth leaf, but then decreased; the exception was for Lp/Wp, which was the reverse. Leaf order of the largest one moved from first position to third as either B or H increased in a community.     

Aboveground and Belowground Impacts Following Removal of the Invasive Species Baby's Breath (Gypsophila paniculata) on Lake Michigan Sand Dunes

The removal of invasive species is often one of the first steps in restoring degraded habitats. However, studies evaluating effectiveness of invasive species removal are often limited in spatial and temporal scale, and lack evaluation of both aboveground and belowground effects on diversity and key processes. In this study, we present results of a large 3‐year removal effort of the invasive species, Gypsophila paniculata, on sand dunes in northwest Michigan (USA). We measured G. paniculata abundance, plant species richness, plant community diversity, non‐native plant cover, abundance of Cirsium pitcheri (a federally threatened species endemic to this habitat), sand movement, arbuscular mycorrhizal spore abundance, and soil nutrients in fifteen 1000 m² plots yearly from 2007 to 2010 in order to evaluate the effectiveness of manual removal of this species on dune restoration. Gypsophila paniculata cover was greatly reduced by management, but was not entirely eliminated from the area. Removal of G. paniculata shifted plant community composition to more closely resemble target reference plant communities but had no effect on total plant diversity, C. pitcheri abundance, or other non‐native plant cover. Soil properties were generally unaffected by G. paniculata invasion or removal. The outlook is good for this restoration, as other non‐native species do not appear to be staging a “secondary” invasion of this habitat. However, the successional nature of sand dunes means that they are already highly invasible, stressing the need for regular monitoring to ensure that restoration progresses.     

Responses of Hedysarum Laeve, a guerrilla clonal semi-shrub in the Mu Us sandland, to local sand burial

In arid and semi-arid inland deserts,one of the environmental stresses for plants is recurrent sand burial,which can influence the physical and biotic microenvironments of the plants and soil.Previous studies have shown that different levels of sand burial have different effects on plants.Slight sand burial could increase the height increment,leaf biomass and the number of new ramets of the plants while heavy sand burial could impair the growth of the plants and even decrease their chances of survival.In other words,below a certain threshold level of burial,the growth of plants is stimulated probably because of multiple factors.However,as the level of burial increases,the positive response starts to decline until it becomes a negative value.Arid and semi-arid inland deserts are frequently colonized and stabilized by many rhizomatous clonal plants.Clonal physiological integration often helps clonal plants buffer local environmental stress encountered by ramets.A rhizomatous clonal semishrub,Hedysarum laeve (H.laeve),is the dominant plant species and important for vegetation restoration in the Mu Us sandland.To investigate whether clonal integration can increase the threshold of sand burial and help rhizomatous H.laeve tolerate heavy sand burial,we conducted a field experiment.The results showed that slight sand burial could accelerate ramet growth and enhance leaf biomass,stem biomass and shoot biomass,while heavy sand burial reducesed the biomass of the plant and impairs survival and growth of the ramets.Clonal integration increased the threshold of sand burial.Under heavy sand burial,ramets connected to other ramets not buried in sand were more in terms of height increment,stem biomass,leaf biomass and shoot biomass compared to the ramets encountering sand burial but disconnected from other ramets.It suggested that clonal physiological integration could help H.laeve ramets tolerate relatively heavy sand burial.We also discussed that clonal integration plays a role in H.laeve presence in the Mu Us sandland.     

Effects of denudation and burial on growth and reproduction of <Emphasis Type="Italic">Artemisia ordosica</Emphasis> in Mu Us sandland

Plants in a dune environment have a high risk of being denudated by wind or buried by sand. We conducted a field experiment to assess growth and reproductive performance as well as biomass allocation of Artemisia ordosica after denudation and burial. Height growth decreased after denudation, but remained constant after burial; biomass was reduced by severe burial, but not by moderate ones. Vegetative growth measured by current-year generated vegetative branches declined with increasing severity of burial. Reproductive maturity was not affected by either denudation or burial, and reproduction decreased only after severe denudation. After denudation, biomass allocated to reproduction was greatly reduced, as shown by strongly reduced biomass allocation to fruits and whole reproductive branches, but that allocated to current-year vegetative growth was maintained. Biomass allocation to reproduction and vegetative growth remained rather stable after burial. Current-year reproductive-to-vegetative-biomass ratio was lower after denudation than after burial, indicating that vegetative growth was more favored by A. ordosica after denudation. These modifications are adaptive in the fluctuating environment of mobile sand dunes where denudation and burial frequently occur.     

小叶锦鸡儿幼苗对沙埋的生态适应和生理响应

以中国半干旱地区固定、半固定沙地分布最广泛的灌木种之一小叶锦鸡儿(Caragana microphylla)为对象,于2010～2011年在内蒙古科尔沁沙地测定了不同深度沙埋下其幼苗的存活率、株高、渗透调节物质含量、保护酶活性和膜透性变化,探讨小叶锦鸡儿对沙埋的生理响应特征。结果显示:(1)与对照相比,当沙埋深度为小叶锦鸡儿幼苗株高的25%～75%时,存活率差异不显著;当沙埋深度为50%～75%时其存活率增加,但株高明显下降,沙埋对其生长具有抑制作用;当沙埋达到其株高100%时幼苗全部死亡。(2)不同沙埋深度对小叶锦鸡儿幼苗叶片细胞膜透性无显著影响,对沙埋深度为其株高25%时,叶片含水量增加,MDA含量下降,POD活性增强,细胞膜未受到伤害;沙埋深度为其株高50%～75%时,叶片含水量下降,MDA含量增加,但SOD和POD对细胞膜起到了协同保护作用,细胞膜也未受到损伤;沙埋深度为株高25%～75%情况下,细胞膜未受到损伤,可溶性糖和脯氨酸含量变化不明显。研究认为:小叶锦鸡儿幼苗只能耐受部分沙埋,完全沙埋则会导致其幼苗全部死亡,沙埋胁迫下可溶性糖和脯氨酸等渗透调节物质含量反应迟缓可能是其耐沙埋能力较弱的主要生理原因之一。     

The role of mycorrhizal fungi in growth enhancement of dune plants following burial in sand

1. Burial in sand of Agropyron psammophilum and Panicum virgatum plants had a stimulating effect on carbon dioxide exchange rate, leaf area and biomass, irrespective of whether sand used for burial did or did not contain mycorrhizal fungi.
2. Plants of both A. psammophilum and P. virgatum species grown in mycorrhiza-containing sand and then buried with mycorrhiza-containing sand had the highest CO₂ exchange rate, leaf area and biomass.
3. The growth stimulation following a burial episode is probably a composite response of several factors. The major contribution of mycorrhizal fungi will possibly be the exploitation of resources in the burial deposit.     

Clonal integration enhances survival and performance of Potentilla anserina, suffering from partial sand burial on Ordos plateau, China

On Ordos plateau, a semi-arid, desertified area in China, sand burial is a common stress factor for plants. The extent to which sand burial occurs is heterogeneous and unpredictable in space and in time. Therefore, clonal fragments (i.e., interconnected ramets of a clonal plant) often experience partial sand burial, with some ramets buried in sand while the rest may remain unburied. It was hypothesized that clonal fragments are able to benefit from clonal integration, in case they experience partial sand burial. A pot experiment was conducted with Potentilla anserina, a stoloniferous herb often found on Ordos plateau. We used clonal fragments consisting of four interconnected ramets. In the experiment, the two proximal (older) ramets were unburied while the two distal (younger) ramets were either unburied (control) or buried with a 2, 4 or 6 cm deep layer of sand (burial treatments). The stolon connection between the proximal and the distal ramets was either severed or left intact. Stolon severing dramatically decreased the survival of buried ramets. Stolon severing and sand burial had significant effects on plant performance in terms of biomass production, number of leaves and leaf area. A cost–benefit analysis based on performance measures shows that the proximal ramets supported their connected distal ramets and did not incur any cost from this resource export. These results suggest that clonal integration, which is one of the functionally most important consequences of clonal growth, contributes significantly to our test species' capacity to withstand partial sand burial on Ordos plateau, a semi-arid and desertified area of China.     

Consequences of variation in species diversity in a community of root-feeding herbivores for nematode dynamics and host plant biomass

To date, no study has explicitly addressed effects of variation in species diversity of root‐feeding herbivores on host plant biomass. Root‐feeding nematodes typically occur in multi‐species communities. In a three‐year field experiment, we investigated how variation in species diversity of root‐feeding nematodes affected nematode dynamics and response of the dune grass Ammophila arenaria to root‐feeder activity. This plant species needs regular burial by fresh beach sand to remain vigorous, suggesting that A. arenaria benefits from a temporary escape from root‐feeding soil organisms and that root‐feeders are involved in plant degeneration in stabilized dunes. We created series of ceased and continued sand burial and added the endoparasitic nematodes Meloidogyne maritima, Heterodera arenaria and Pratylenchus penetrans alone or in combination to A. arenaria. We included treatments with and without the whole soil community, measured plant biomass and quantified numbers of nematodes. Addition of H. arenaria and P. penetrans decreased numbers of M. maritima juveniles and delayed the first appearance in time of both juveniles and females, while numbers of males only decreased when plants had been buried. Burial with sand and addition of the other two endoparasites affected numbers of H. arenaria juveniles, while numbers of P. penetrans were low and not affected. Shoot biomass of A. arenaria was lower when M. maritima had been added alone than when the three species had been added together. Addition of root zone soil decreased biomass of all plant parts. Burial with sand decreased aboveground shoot biomass, whereas it increased belowground shoot and root biomass. Our results point at idiosyncratic effects of nematode diversity on A. arenaria biomass. Heterodera arenaria and P. penetrans protected their host by reducing numbers and delaying activity of M. maritima to a later stage in the growth season, when root‐feeding activity was less harmful for plant biomass development.     

沙埋对中间锦鸡儿幼苗生长发育的影响

在沙地环境中,沙埋是影响植物幼苗存活的一个重要因素。中间锦鸡儿(C arag ana interm ed ia)是浑善达克沙地和毛乌素沙地固定和半固定沙丘上常见的一种沙生灌木。为了研究沙埋对其幼苗存活以及生长的影响,对出土后生长了1个月的中间锦鸡儿实生苗进行沙埋实验,实验处理:TC为对照,T1为沙埋幼苗地上部的1/3,T2为沙埋幼苗地上部的1/2,T3为沙埋到幼苗的近顶端。实验持续4周,每周收获1次。结果表明:TC、T1和T2没有幼苗死亡,T3有20%的幼苗死亡;沙埋对中间锦鸡儿幼苗的生物量、根冠比、根生物量比和茎生物量比有显著影响,叶片生物量比各处理没有显著差异,沙埋对幼苗的叶面积没有显著影响,在第1周T2和T3的小叶片面积显著大于TC,在其余3周,TC与T1、T2的小叶片面积没有显著差异,与全埋则有显著差异;沙埋对幼苗的相对生长速率(RGR)和净同化速率(N AR)产生显著影响,实验结束时,T3处理幼苗的RGR和N AR显著地小于其它3个处理。上述结果表明,部分沙埋(T1、T2)对沙生灌木中间锦鸡儿幼苗的生长发育没有显著的影响,而全部沙埋(T3)对其幼苗的生长发育产生显著的抑制,中间锦鸡儿是一个比较耐沙埋的物种。     

海滨沙地单叶蔓荆匍匐茎对沙埋适应的生长对策

单叶蔓荆(Vitex trifolia var.simplicifoli)是一种耐盐、耐旱固沙地被植物。依据海滨沙地自然沙埋特点对单叶蔓荆匍匐茎进行了不同厚度(半埋和全埋)和不同长度交叉沙埋处理,研究探讨了单叶蔓荆沙埋适应生长对策,为其开发利用、科学管理和海滨环境修复提供指导。结果表明,正常情况下,单叶蔓荆匍匐茎基部和中部生长缓慢,顶部生长快。轻度(沙埋匍匐茎基部)和中度(沙埋匍匐茎基部和中部)半埋和全埋使匍匐茎顶部生长加速,茎长增长量较对照高出1.5到3.1倍;但重度(沙埋整个匍匐茎)半埋和全埋使匍匐茎顶部净增长量减少12%和13%。在20d沙埋中,对照整个匍匐茎各段均无不定根长出,但不同程度半埋和全埋沙埋处理下沙下匍匐茎上均长出不定根,重度半埋使不定根生长受抑;同时匍匐茎上各段茎生物量上升,枝叶生物量下降,且随着沙埋程度的增加而增减幅度提高,在重度半埋和全埋达到最大。在轻度和中度半埋和全埋下,匍匐茎上未沙埋部位枝条生长加速。研究表明,在自然环境中,单叶蔓荆匍匐茎顶端是一个对环境变化反应敏感的部位,并与沙埋后单叶蔓荆茎延伸生长和植株能否生存密切相关。当匍匐茎顶部没被沙埋时,沙埋促进沙埋部位匍匐茎和枝叶中物质转移,加速匍匐茎顶部快速生长和物质积累以弥补沙埋带来的损伤维持物质和能量的代谢平衡。沙埋后,单叶蔓荆以茎顶端快速生长、形成不定根、枝条生长维持茎水分平衡和能量和物质代谢平衡,以快速生长摆脱沙埋影响的生长方式为其对沙埋环境的重要适应对策。因此,在海岸沙地单叶蔓荆种群管理和维护中,在强风移沙引起的重度沙埋后,及时剥离匍匐茎顶部沙子对维护单叶蔓荆种群的延续生存和扩散均有重要作用。     

Biomass reallocation and the mobilization of leaf resources support dune plant growth after sand burial

The stimulation of dune plant growth in response to burial is a vital attribute allowing survival in areas of mobile sand. Numerous resource-related and physiological mechanisms of growth stimulation have been suggested in the past, but few have been tested comparatively. Manipulation experiments using Scaevola plumieri, an important subtropical coastal dune forming species, demonstrated that physiological shifts were of great importance in determining the nature of the stimulation response to burial. The production of stem length and replacement of leaf area were stimulated by burial, whereas net mass production was similar between buried and unburied treatments. Remobilization of buried leaf resources, seasonal effects, and a shift in biomass allocation to stem production played the greatest role in the compensatory growth response. Other factors, such as increased soil nutrients, changes in photosynthesis, and changes in the costs of producing tissue were of less importance. Thus, the stimulated growth of species adapted to live on mobile dunes is explained by a number of resource-related and physiological mechanisms acting in concert.     

The role of ethylene and darkness in accelerated shoot elongation of Ammophila breviligulata upon sand burial

The impact of elevated ethylene concentrations and darkness on the growth and development of shoot organs of Ammophila breviligulata was investigated under experimental conditions in a complete two-way design. The results were compared with data of partially sand buried plants. Enhanced ethylene concentrations and sand burial stimulated the formation of new stem nodes, a prerequisite for burial-induced shoot elongation. However, internode elongation itself could not be promoted by the phytohormone ethylene, by darkness, or by their interaction. Sand burial inhibited the formation of rhizomes and tillers and the investment in root and rhizome biomass. Darkness mimicked this effect for the number of rhizomes and the biomass allocated to roots and rhizomes, indicating that the change in light regime upon sand burial may play an important role in the signal transduction chain that leads to a different allocation pattern in A. breviligulata. The results are discussed within the context of alternative signals that might initiate the internode elongation response in sand-buried A. breviligulata plants. Received: 5 July 1997 / Accepted: 28 February 1998     

毛乌素沙地游击型克隆半灌木羊柴对局部沙埋的反应

不同程度的沙埋是生长在干旱和半干旱区内陆沙丘的植物经常遭遇的事件,沙埋可以改变植物所处的生物和非生物环境条件。已有研究表明不同程度的沙埋对于植物的影响不同。轻微程度的沙埋可以增加植物高度、促进生物量的积累和新生分株的产生。如果沙埋强度不断增加,对植物的影响由正效应逐渐转变为负效应。即超过一定沙埋阈值后,沙埋会削弱植物的生长,甚至影响植物的存活。干旱和半干旱区内陆沙丘中常常生长着许多克隆植物,克隆整合常常可以缓解克隆植物分株所遭受的局部环境胁迫。根茎型克隆植物羊柴(Hedysarum laeve)是毛乌素沙地的优势半灌木之一,也是当地重要的固沙植物。为了探讨克隆整合的作用是否可以提高沙埋阈值,并有助于羊柴忍受高强度的沙埋,以其为研究对象开展了野外实验。结果表明:轻微程度的沙埋(例如沙埋深度是原始羊柴分株高的10%~20%)可以加速羊柴分株的高生长,提高叶片生物量、茎生物量以及整个地上部分的生物量。高强度的沙埋(例如沙埋深度是原始羊柴分株高的80%~100%)会削弱羊柴分株的存活和生长。在与不遭受沙埋分株相连的情况下,羊柴分株遭受沙埋的阈值高于没有分株相连的,而且在高强度的沙埋下,前者(有分株相连的遭受沙埋的分株)比后者(没有分株相连的遭受沙埋的分株)在株高增量、茎生物量、叶片生物量以及地上分株生物量上都要显著高。这暗示着克隆整合提高了羊柴遭受沙埋的阈值并有助于羊柴分株忍受高强度的沙埋。     

克隆植物沙拐枣的母株和分株对风蚀沙埋的生理生态响应

以中国荒漠区优良的防风固沙克隆灌木沙拐枣为对象,研究了长期风蚀、沙埋环境下沙拐枣母株和克隆分株的同化枝对环境异质性的响应。结果发现:(1)风蚀母株、风蚀分株的叶绿素含量、净光合速率、气孔导度、蒸腾速率、胞间CO2浓度和水分利用效率只有沙埋分株的一半左右,导致同化枝的长度、数量、簇数也仅是沙埋分株的一半,而且风蚀母株的果实宽和果实长也都最小,但浅沙埋有利于沙拐枣的生长和繁殖,表明严峻的风蚀对母株和分株的生长与繁殖都产生了胁迫,但浅沙埋有利于沙拐枣的生长和繁殖。(2)风蚀母株倒伏后同化枝的形态特征是基部优于中部优于顶部,表明严峻风蚀下母株的死亡是从顶部-中部-底部逐渐舍弃的过程。(3)母株的全部根系以及风蚀水平根全部裸露在外但依然能够存活,间接证明沙拐枣克隆整合的方向性——不仅可在分株间进行传递,分株-母株间也可进行传递,否则遭受严峻风蚀胁迫的母株和克隆分株会直接死亡。本研究结果为沙拐枣克隆生长对风沙环境的生态适应机制提供了基础,也是对植物克隆生态学在自然异质环境中研究缺乏的有效补充。