Effects of flooding and drought on stomatal activity, transpiration, photosynthesis, water potential and water channel activity in strawberry stolons and leaves

Transpiration, xylem water potential and water channel activity were studied in developing stolons and leaves of strawberry (Fragaria × ananassa Duch.) subjected to drought or flooding, together with morphological studies of their stomata and other surface structures. Stolons had 0.12 stomata mm^–2 and a transpiration rate of 0.6 mmol H₂O m^–2 s^–1, while the leaves had 300 stomata mm^–2 and a transpiration rate of 5.6 mmol H₂O m^–2 s^–1. Midday water potentials of stolons were always less negative than in leaves enabling nutrient ion and water transport via or to the strawberry stolons. Drought stress, but not flooding, decreased stolon and leaf water potential from –0.7 to –1 MPa and from –1 to –2 MPa, respectively, with a concomitant reduction in stomatal conductance from 75 to 30 mmol H₂O m^–2 s^–1. However, leaf water potentials remained unchanged after flooding. Similarly, membrane vesicles derived from stolons of flooded strawberry plants showed no change in water channel activity. In these stolons, turgor may be preserved by maintaining root pressure, an electrochemical and ion gradient and xylem differentiation, assuming water channels remain open. By contrast, water channel activity was reduced in stolons of drought stressed strawberry plants. In every case, the effect of flooding on water relations of strawberry stolons and leaves was less pronounced than that of drought which cannot be explained by increased ABA. Stomatal closure under drought could be attributed to increased delivery of ABA from roots to the leaves. However, stomata closed more rapidly in leaves of flooded strawberry despite ABA delivery from the roots in the xylem to the leaves being strongly depressed. This stomatal closure under flooding may be due to release of stress ethylene. In the relative absence of stomata from the stolons, cellular (apoplastic) water transport in strawberry stolons was primarily driven by water channel activity with a gradient from the tip of the stolon to the base, concomitant with xylem differentiation and decreased water transport potential from the stolon tip to its base. Reduced water potential in the stolons under drought are discussed with respect to reduced putative water channel activity.     

Influence of Ethylene and Kinetin on Tuberization and Enzyme Activity in Solanum tuberosum L.: Stolons Cultured in vitro

The response of potato stolons, cultured in vitro, to ethylenewas investigated utilizing 2-chloroethyl-phosphonic acid asthe source of ethylene. Concentrations of 0.067, 0.67, 6.7,and 67 µm did not promote tuber formation which occurredin the presence of 16 µm kinetin. In the presence of 2-chloroethyl-phosphonicacid stolon branching was promoted and they maintained a diageotropicgrowth habit in contrast to the negatively geotropic growthhabit of kinetin-treated cultures. The amount of soluble sugars accumulating at the stolon tipwas similar in the presence of kinetin and 2-chloroethylphosphonicacid. However, the level decreased in kinetin-treated stolonsas starch synthesis increased whereas a high specific activitywas maintained in the presence of 2-chloroethylphosphonic acidwhere no starch synthesis occurred. Higher levels of sucroseoccurred in kinetin-treated cultures. Both 2-chloroethylphosphonic acid and kinetin decreased invertaseactivity at the stolon tip. Peroxidase activity increased withtime in response to 2-chloroethylphosphonic acid whereas activityincreased appreciably in kinetin-treated stolons only afterday 5. Generally higher levels of 3'nucleotidase activity existedin 2-chloroethylphosphonic acid-treated stolons. RNase activitydecreased in kinetin treated stolons while activity increasedin the presence of 2-chloroethylphosphonic acid. It is suggestedthat ethylene may influence stolon growth but may not be directlyinvolved in tuber initiation.     

Ditylenchus dipsaci Infestation of Trifolium repens. II. Dynamics of Infestation Development

Trifolium repens (white clover) stolons were inoculated with Ditylenchus dipsaci (stem nematode), and the development of resulting infestations was monitored. Nematodes initially remained confined to superficial locations, concentrating in petiole axils near inoculation points. They were able to migrate slowly from the inidal inoculation points and infest adjacent axils, especially in regions near the stolon tip. As time progressed, in some axils, nematodes migrated through the stolon epidermis and colonized slowly expanding subepidermal pockets of host tissue (ca. 0.2-mm length of stolon/day). In these loci nematodes established exponentially increasing populations, but the rates of locus expansion remained constant, indicating that locus expansion was limited by unidentified host-dependent factors. As a result of increasing population pressure within subepidermal loci, J4 entered a "diapause" state and the rate of egg production by adults declined, thereby reducing rate of population growth to more sustainable levels. Typically, these populations peaked at ca. 10,000 individuals in ca. 160 days occupying 3-cm lengths of stolon. Thereafter, heavily infested regions of stolons started to die, leading to the formation of longitudinal splits in their epidermis. In other axils, nematodes did not migrate into the stolons but remained confined to axils. Some of these populations increased a hundred-fold in 95 days, with population growth ending when petioles started to die. Host plant stolon morphology was affected only when subepidermal stolon populations developed high population levels (>100 nematodes) within close proximity (<2 cm) to active terminal meristems. This occurred either when axillary buds became active on previously infested nodes or when nematodes established endoparasitic populations at locations near the stolon tip during winter and spring, when the rate of stolon extension was limited by low light intensity. Affected stolon tips could "escape" from the influence of such infestations when light intensity and temperature increased. Nematode activity was limited by low temperature rather than light intensity. Global warming is likely to lead to greater damage to infested plants during the winter and early spring because the predicted milder winter temperatures will enhance nematode activity but not necessarily promote stolon growth.     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation     

The Role of Gibberellin, Abscisic Acid, and Sucrose in the Regulation of Potato Tuber Formation in Vitro

The effects of plant hormones and sucrose (Suc) on potato (Solanum tuberosum L.) tuberization were studied using in vitro cultured single-node cuttings. Tuber-inducing (high Suc) and -noninducing (low Suc or high Suc plus gibberellin [GA]) media were tested. Tuberization frequencies, tuber widths, and stolon lengths were measured during successive stages of development. Endogenous GAs and abscisic acid (ABA) were identified and quantified by high-performance liquid chromatography and gas chromatography-mass spectrometry. Exogenous GA_4/7 promoted stolon elongation and inhibited tuber formation, whereas exogenous ABA stimulated tuberization and reduced stolon length. Indoleacetic acid-containing media severely inhibited elongation of stolons and smaller sessile tubers were formed. Exogenous cytokinins did not affect stolon elongation and tuber formation. Endogenous GA₁ level was high during stolon elongation and decreased when stolon tips started to swell under inducing conditions, whereas it remained high under noninducing conditions. GA₁ levels were negatively correlated with Suc concentration in the medium. We conclude that GA₁ is likely to be the active GA during tuber formation. Endogenous ABA levels decreased during stolon and tuber development, and ABA levels were similar under inducing and noninducing conditions. Our results indicate that GA is a dominant regulator in tuber formation: ABA stimulates tuberization by counteracting GA, and Suc regulates tuber formation by influencing GA levels.     

Effect of Carbohydrate Demand on the Remobilization of Starch in Stolons and Roots of White Clover (Trifolium repens L.) after Defoliation

White clover plants were grown from stolon tips in growth cabinetsand then defoliated. Thereafter, changes in the contents ofnon-structural carbohydrates such as starch, sucrose, glucose,fructose, maltose, and pinitol in stolons and roots were monitored.Initial contents of carbohydrate reserves, photosynthetic supplyof new carbohydrates and carbohydrate demand after defoliationwere varied by growing the plants at various CO₂ partial pressures,by varying the extent of defoliation and by removing eitherroots or stolon tips at the time of defoliation. Remobilization of carbohydrate reserves in stolons increasedproportionally to their initial contents and was greater whenplants had been severely defoliated, suggesting that carbohydrateswere remobilized according to availability and demand. Starchwas the predominant reserve carbohydrate. Starch degradationwas associated with decreased contents of sucrose, glucose andfructose in young stolon parts and roots but not in old stolonparts suggesting that starch degradation was not strictly controlledby the contents of these sugars. A decrease in the demand forcarbohydrates by removal of roots did not decrease starch degradationbut increased the contents of sucrose, glucose, and fructose.Removal of stolon tips decreased starch degradation and contentsof sucrose, glucose, and fructose. The results suggest thatstarch degradation was controlled by a factor other than sucrose,glucose, and fructose which was exported from stolon tips, e.g.gibberellin. Key words: White clover, storage carbohydrates, remobilization, regrowth     

Senescence and decomposition of white clover stolons in grazed upland grass/clover swards

The fate of marked sections of stolons of white clover (Trifolium repens) over a 50-week period from May 1987 was followed in grazed grass/clover swards maintained at 5-cm sward surface height with and without N fertiliser. There was little effect of N treatment on the pattern of survival of stolon sections. The proportion of live stolons recovered decreased during the experiment, and in May 1988 on average only 29% of the marked sections remained alive. At all harvests only a small percentage of stolon sections showed signs of senescence; the maximum percentage, on average 20% of those marked, occurred in autumn, 15–20 weeks after marking. Following this period, i.e. in late autumn/winter, the most rapid increase in percentage of decomposed stolons was measured. Over 50% of stolon sections were buried within the 5-week period following marking and nearly all were buried after 20 weeks; generally a much smaller proportion of stolon tips was buried. Nutrient concentrations of N, P and K fell to their lowest levels in autumn, before increasing in the following spring. Results are discussed in relation to the cycling of nutrients via stolon senescence.     

Contribution from Stolons and Roots to Estimates of the Total Amount of N2Fixed by White Clover (Trifolium repensL.)

Growth and N-accumulation rates in leaves, stolons and rootsof individual white clover plants were studied in three experimentsusing two methods. In a growth chamber experiment, the relativedifferences between tissues were found to be almost constantfor a wide range of clover plant sizes. The stolon dry matter(DM) production was 56% and the root DM production 40% of theDM production in leaves. The N yield of stolons was 30% whileN yield in roots was 34% of N yield in leaves. The effect ofN application on these relations was investigated in a glasshouseexperiment. Application of N reduced the root:shoot N ratiofrom 0.50 to 0.28, whereas the stolon+root:leaf N ratio (i.e.for abovevs.below cutting-height tissues) was only reduced from0.97 to 0.80. In a field trial with two contrasting N regimes,growth and N accumulation were measured on individual cloverplants. Dinitrogen fixation was estimated by¹⁵N isotope dilutionbased on analysis of leaves-only or by including stolons. Usingleaves-only did not affect the calculation of percentage ofclover N derived from N₂fixation (% Ndfa) since the¹⁵N enrichmentwas found to be uniform in all parts of the clover. A correctionfactor of 1.7 to account for N in below cutting-height tissueis suggested when N₂fixation in white clover is estimated byharvesting the leaves only.Copyright 1997 Annals of Botany Company Leaves; N accumulation; N₂fixation; ¹⁵N isotope dilution; pastures; roots; root/shoot ratio; stolons; Trifolium repensL.; white clover     

Water Relations of White Clover (Trifolium repens): Water Potential Gradients and Plant Morphology

Water potential, osmotic potential, pressure potential and relativewater content were measured in stolons and leaves of white cloverplants grown under a range of conditions of water supply andevaporative demand. The importance of adventitious roots fromthe nodes was examined. Gradients along stolons were alwaysextremely small, of the order of only 01 MPa. Stolon up waterpotential was representative of plant water status regardlessof stolon length, presence/absence of nodal roots, degree ofwater stress and evaporative demand. It is concluded that waterconduction along stolons was very good. Gradients were foundto exist along petioles; they may have a greater resistanceto water flow than stolons. The relationship between water fluxand stem anatomy, and the importance of differential flow ratesthrough stolons and petioles to plant behaviour during waterstress, are discussed. Trifolium repens L., white clover, water relations     

The effect of moisture stress on stolon and adventitious root development in white clover (Trifolium repens L.)

Summary Humidity, at the young nodes of white clover stolones, varied by enclosing nodes in the atmosphere above a range of saturated solutions, inhibited root initiation at 85% RH or less. The threshold humidity for root initiation increased to about 93% on young nodes subject to moisture stress or old nodes on well watered plants in which root initiation had been previously suppressed by low humidity.Roots at old nodes and at the three youngest on stolons were either subject to moisture stress or adequately watered. Growth of young roots and N₂-fixation were more adversely affected by the direct effects of drought than by subjecting old roots to drought. Although old roots under stress affected new root growth and N₂-fixation, length of roots and lateral root number were little affected. By contrast stolon growth was affected more by stress to old roots than to young nodes, although after 6 weeks the contribution made by young roots to stolon growth was almost as high as old roots.The data suggest that deep roots at old nodes will allow clover stolons to grow during drought due to the high acropetal movement of water but initiation of roots and functioning of young roots at the soil surface will be adversely affected, with possible implications on the persistence of clover.     

Phosphate Uptake and Transport by Roots and Stolons of Intact White Clover (Trifolium repens L.)

The extent to which phosphate can be absorbed directly fromthe outer medium by stolon internodes and contribute to thetotal accumulation of phosphate by intact plants of white clover(Trifolium repens L. cv. Blanca) was assessed in hydroponicexperiments in a controlled environment room. The uptake ofphosphate by intact roots or stolons was measured by sealinga segment (6-0 mm long) across a flow-cell in which ³²P-labellednutrient solution was circulated for 24 h, the rest of the rootsystem receiving unlabelled nutrient solution. The rate of uptakeof phosphate (µmol g^–1 d^–1 dry wt. basis)by roots was more than 300 times that by intact stolons. Pretreatmentof stolons by gentle abrasion to remove cuticle, so as to simulatethe condition of stolons in the field, increased the uptakeof phosphate 7-fold compared with that of intact stolons. However,the potential of stolons to contribute to the P status of whitedover in the field was calculated to be small (5%). When an incision was made through the hypodermal layer of stolons,the rate of phosphate uptake greatly increased, attaining 71%of that by root segments. This increase, which was greater athigher phosphate concentrations, indicates that the suberi.zedhypodermis constitutes a major barrier to the influx of phosphatein the stolon. After withholding phosphate for different time intervals, thesubsequent rate of phosphate uptake by roots was increased 2-3-foldafter 2 d phosphate deprivation and 3-4-fold after 6 d or 13d phosphate deprivation. A higher proportion of absorbed phosphatewas transported to shoots in phosphate-deprived plants. After1 d of uptake following restoration of the phosphate supply,the concentrations of labelled phosphate in shoots were greaterthan in control plants, although the concentrations of labelin roots was less. However, the rate of uptake of phosphateby stolons, following deprivation, was not significantly increased.These results suggest that the mechanism regulating the enhancedrate of phosphate loading into the xylem, initiated by a periodof phosphate deprivation, is specific to roots and is not inducedin stolons. The results are discussed in relation to the growth and acquisitionof phosphate by white clover in the field. Key words: Nutrient deficiency, phosphate, stolons, transport (ions), Trifolium repens     

Role of Ethylene in the Geotropic Response of Bermudagrass (Cynodon dactylon L. Pers.) Stolons

We studied the relationship between ethylene and gravity-induced upward bending of bermudagrass (Cynodon dactylon L. Pers.) stolons. Ethylene production begins within 3 hours of the onset of gravistimulation, and increases thereafter until the 15th hour, after which it declines. There is a close positive relationship between ethylene production and upward bending during the first 12 hours of gravistimulation. Incubation of stolons with AgNO₃ did not prevent ethylene evolution but delayed upward bending. In addition, ethylene production was 10-fold greater and peaked earlier in gravistimulated nodes incubated with 1-aminocyclopropane 1-carboxylic acid. The gravitational stimulation could be due to an increase in both 1-aminocyclopropane 1-carboxylic acid synthase and the ethylene forming enzyme. The results suggest that ethylene promotes the activity of indoleacetic acid.     

Synchronization of tuber formation in potato grown in vitro by cell division synchronization in axillary meristems of stem explants

Optimization of in vitro tuberization (formation and growth of stolons and microtubers) by synchronization of cell divisions in axillary meristems of initial stem explants induced by low nonfreezing temperatures was studied in potato (Solanum tuberosum L., cv. Lugovskoi) plants. The proportion of simultaneously dividing cells in axillary meristems of stem explants subjected to 2-h cold treatment at 4°C was in 2.6 times greater than in control material (without chilling). The analysis of growth of stolons and microtubers produced from the explants exposed to cold showed that synchronization of cell divisions in the meristems of initial explants resulted in synchronization of stolon and microtuber formation and production of microtubers of identical physiological age.     

Axillary Bud Development in White Clover in Relation to Defoliation and Shading Treatments

A period of growth under shade netting in the glasshouse allowedthe cultivation of white clover stolons with an accumulationof undeveloped axillary buds similar to that often found onstolons from grass/clover swards. The subsequent capacity ofthese nodes to develop branches under different circumstanceswas investigated in three experiments. Removal of the laminaeand petioles subtending sets of four buds along a stolon reducedthe rate at which branches were initiated from the buds. Treatmentsin which petioles, or petioles plus laminae, were retained initiatedbranches more quickly. Shading the stolons reduced both therate of initiation and the percentage of buds which developed,unless both petioles and laminae were retained. There was someevidence that conditions applied to individual buds may actin the same way as the same conditions applied to sets of fourbuds and that illuminated nodes may depress the performanceof neighbouring shaded notes. Fewer buds developed at older nodes than at younger nodes duringthe summer, but during the autumn younger buds initially developedmore slowly than older buds. This suggests that buds can developat a younger nodal age in summer than in winter. When leafless stolons were cut up into component internodesbuds developed faster than on intact stolons, provided the budwas located at the end of the internode nearest the main stolongrowing point. If the bud was at the other end, branch developmentwas slower than on intact stolons. The results are discussedin relation to clover growth in sward conditions. White clover, Trifolium repens, axillary bud development, branching, growing points, defoliation, shading     

Xylem and Phloem Transport of Mineral Nutrients from Solanum tuberosum Roots

The xylem and phloem transport of mineral elements from stemnodal roots to the stem and stolon of growing potato (Solanumtuberosum L. cv. ‘Russet Burbank’) plants was investigated.Adventitious roots, originating from below-ground nodes of thestem of potato seedlings, were exposed to solutions of SrCI₂or MnSO₄. Relative elemental concentrations were measured inthe conductive tissues using energy dispersive X-ray analysis.After a 5 h daylight uptake period, Sr (a Ca-transport analogue)levels were elevated in the stem xylem tissue, but Sr did notincrease in the stem phloem, nor was it present in either ofthe conductive tissues of stolons located 1–2 nodes abovethe treated roots. In contrast, elevated levels of Cl, S, andMn were found in stolon xylem and phloem tissue during the sameperiod. The absence of Sr in the stolon after 5 h suggests thatno xylem flow into the stolon occurred during the uptake periodand, furthermore, phloem flow is responsible for the transportof the Cl, S, and Mn into the stolon. Elevated levels of thesemobile nutrients in the xylem of the stolon were attributedto xylem-to-phloem transfer in the stem or leaves, transportto the stolon in the phloem, and phloem-to-xylem transfer inthe stolon. During a 19 h uptake period, some Sr was observedin the phloem tissue of the stem, demonstrating slow exchangeof Sr with sieve elements or proximal phloem parenchyma andcompanion cells. Key words: Calcium, manganese, X-ray analysis     

Nutrient Distribution and Absorption in the Colonial Hydroid Podocoryna carnea Is Sequentially Diffusive and Directional

The distribution and absorption of ingested protein was characterized within a colony of Podocoryna carnea when a single polyp was fed. Observations were conducted at multiple spatial and temporal scales at three different stages of colony ontogeny with an artificial food item containing Texas Red conjugated albumin. Food pellets were digested and all tracer absorbed by digestive cells within the first 2–3 hours post-feeding. The preponderance of the label was located in the fed polyp and in a transport-induced diffusion pattern surrounding the fed polyp. After 6 hours post-feeding particulates re-appeared in the gastrovascular system and their absorption increased the area over which the nutrients were distributed, albeit still in a pattern that was centered on the fed polyp. At later intervals, tracer became concentrated in some stolon tips, but not in others, despite the proximity of these stolons either to the fed polyp or to adjacent stolons receiving nutrients. Distribution and absorption of nutrients is sequentially diffusive and directional.     

Long-term Partitioning, Storage and Remobilization of 14C Assimilated by Trifolium repens (cv. Blanca)

The fourth fully expanded leaf on the main stolon of white cloverplants was exposed to ¹⁴CO₂. Thereafter, quantitative and fractionalanalysis of the partitioning, storage and remobilization afterdefoliation of the ¹⁴C labelled assimilate was sequentiallyconducted over a 2- to 3-week period. In undefoliated plants, most ¹⁴C reached its final destinationwithin 24 h of feeding. Forty percent of assimilated ¹⁴C waslost through respiration, while the rest was exported, predominantlyto meristems, but also to roots, stolons and leaves. The ¹⁴Cinitially translocated to meristems was subsequently recoveredin stolon and leaf tissue as the plants matured. Approximately 10% of assimilated ¹⁴C was invested into long-termstorage in roots and stolons. These reserves were remobilizedafter both partial and total defoliation, and a portion of theremobilized ¹⁴C was incorporated into new growth, Partly defoliatedplants regrew more rapidly than totally defoliated plants, butmore ¹⁴C reserve depletion took place in the totally defoliatedtreatment. Reserve depletion took place from both stolons androots, but stolon reserves were preferentially utilized. Bothhigh and low molecular weight storage compounds were involved. Trifolium repens, white clover, assimilate partitioning, storage, remobilization, defoliation     

Senescence in isolated carnation petals : effects of indoleacetic Acid and inhibitors of protein synthesis

Indoleacetic acid induces senescence in isolated carnation (Dianthus caryophyllus, cv. White Sim) petals, increasing the duration and amount of ethylene production. This effect is inhibited by Actinomycin D, an inhibitor of RNA synthesis, and cycloheximide, a translational inhibitor of protein synthesis. The ability of petals to respond to indoleacetic acid appears to be a function of physiological age. Indoleacetic acid is capable of enhancing ethylene evolution and senescence only in specific portions of the petal.     

Relation of Phytochrome-enhanced Geotropic Sensitivity to Ethylene Production

Brief exposure of etiolated pea (Pisum sativum cv. Alaska) seedlings to red light enhances subsequent development of geotropic curvature of the stem. Both this response and inhibition of ethylene production by red light become maximal 8 hours after illumination. Very low concentrations of applied ethylene inhibit development of geotropic curvature, whereas hypobaric treatment enhances geotropic sensitivity by removing endogenous ethylene. Increased geotropic sensitivity after illumination is accompanied by increased lateral migration of ³H-indoleacetic acid in response to gravity, and ethylene inhibits this lateral migration. It is suggested, therefore, that red light-enhanced geotropic sensitivity is caused by increased lateral auxin transport resulting from a reduction in ethylene production after illumination.     

酸雨和采食模拟胁迫下克隆整合对空心莲子草生长的影响

研究表明克隆整合可以显著提升异质环境中克隆植物的生长,然而当克隆植物遭受均质环境压力时,整合对植物生长影响的研究相对较少。本文以典型入侵克隆植物空心莲子草（Alternanthera philoxeroides）为例,研究均质环境压力酸雨和采食模拟胁迫对空心莲子草生长的影响,以及克隆整合在空心莲子草适应不利环境过程中所起的作用。酸雨设3种浓度梯度：pH值3.5 、pH值4.5和 pH值6.5（对照）;采食设3种水平：不去叶、去叶50%和去叶90%;整合水平：匍匐茎切断和连接。结果表明：无论保持或切断匍匐茎的连接,酸雨处理都不影响空心莲子草生物量。当保持匍匐茎连接时,pH值4.5酸雨处理增加了空心莲子草匍匐茎长度和分株数目,因此,低度酸雨可能对空心莲子草生长有一定的促进作用。同样,无论匍匐茎是否被切断,采食处理都显著降低了空心莲子草克隆片段生物量,而显著增加了叶片数目。当切断匍匐茎连接时,采食处理使空心莲子草分株数目显著增加。本文得出的结论是：空心莲子草能较好地适应酸雨和采食的环境压力,当空心莲子草全部克隆分株遭受均质环境胁迫时,克隆整合并不能显著改善它的生长。