Central Die-back of Monoclonal Stands of Reynoutria japonica in an Early Stage of Primary Succession on Mount Fuji

Reynoutria japonica is a common perennial pioneer species onJapanese volcanoes. In a volcanic desert (1500m above sea level)on Mount Fuji (3776m), central Japan, this species forms circularstands (patches). As a patch develops, shoot density decreasesin its centre (‘central die-back’). The centraldie-back has been considered a key process in the early stagesof primary succession, though its mechanism has been unknown. The pattern of patch development, population dynamics of aerialshoots, and growth patterns of below-ground organs were analysedin order to investigate the mechanism of die-back, and the followingtraits are clarified: (1) central die-back areas occur in mostsmall patches (approx. 1m²) without later successional species;(2) shoot characteristics are dependent both on their positionwithin a patch and on patch size; (3) despite the large differencesin shoot density, neither time course of shoot growth nor theirmortality differs between the centre and periphery of patches;and (4) rhizomes ofR. japonicagrow outwards with regular sympodialbranching. From these results, it is concluded that neither interspecificnor intraspecific competition is likely to be a primary causeof the die-back phenomenon, but that central die-back is broughtabout intrinsically by the growth pattern of the rhizome systems.We also discuss the importance of the central die-back in facilitatingestablishment of later successional species in the early stagesof primary succession. Clonal plant; central die-back; competition; facilitation; Japanese knotweed; Mount Fuji; primary succession;Reynoutria japonica ; rhizome growth; volcanic desert     

Nitrogen translocation via rhizome systems in monoclonal stands ofReynoutria japonica in an oligotrophic desert on Mt Fuji: Field experiments

Mechanisms which enableReynoutria japonica, a dominant pioneer herb, to be successful in maintaining large stands in an oligotrophic volcanic desert on Mt Fuji were investigated with special reference to its nitrogen acquisition.Reynoutria japonica forms circular stands, each of which comprises only one genet. As a stand develops outwards, the number of aerial shoots per unit area decreases in the center. Shoots grow vigorously in the peripheral area where the available nitrogen from soil and precipitation (about 2.4 g m⁻² year⁻²) was much less than total nitrogen in the shoots (6.1–9.1 g m⁻²). Leaf nitrogen content per unit mass was also greater in the leaves of the peripheral shoots. When rhizomes extending radially from the center to the periphery were severed, the dry mass of shoots in the periphery diminished by 75% on a ground area basis. In the periphery, leaf nitrogen content also reduced significantly and no flowers were produced. When fertilizer was applied to the peripheral shoots with severed rhizomes, neither growth, survival nor flower production of the shoots was significantly smaller than the control levels. In these shoots, it is also found that the nitrogen content in the youngest leaves decreased for about 1 month and then increased to above that in the control leaves. These results suggest that (i) nitrogen accumulated in the central part is translocated to peripheral shoots via rhizomes, and that long-distance translocation enables the stands to develop outwards, and (ii) aerial shoots in the periphery utilize the nitrogen translocated by rhizomes in the beginning of the growth season, whereas once the shoots have established, they begin to take up nitrogen with their own roots. Since the peripheral shoots are in sunnier environments than the shoots inside the stand, the acropetal nitrogen translocation via rhizomes will raise the production efficiency of a whole stand.     

Phragmites australis at an extreme altitude: Rhizome architecture and its modelling

In central EuropePhragmites australis is a lowland plant, occurring rarely up to the tree line. In the Velká Kotlina cirque (Jeseníky mountains, NE Czech Republic), where it reaches its maximum altitude at about 1350 m a.s.l., its culms are 0.5–0.7 m high and the plants flower only in some years. During the last decade no germinable seeds have been observed. The architecture ofPhragmites rhizomes from this site was studied on seven randomly selected clonal fragments. They consisted of 3 to 10 partial tussocks (clumps) and 4 to 17 green shoots. The total length of the rhizomes was 9.7 to 50 m per plant. The number of nodes per plant was 96 to 431 and the longest internodes were 83 mm long. The number of side branches was 31 to 105 per plant. The branching angle depended on the type of branched rhizome. The mean angles of horizontal rhizomes, which connect individual tussocks, were relatively wide (modus 45°, arithmetic mean 37°), whereas within a tussock much sharper angles of branching prevailed (modal value 5°, arithmetic mean 15°). The mean internode-to-internode angle on continuing rhizomes was about 8°, with a wide variation. An architectural, spatially-explicit model ofPhragmites rhizome growth has been developed, showing that thePhragmites population in the studied locality can be maintained by vegetative multiplication, and seedling recruitment is not needed for its long-term persistence.     

Ultrastructural Investigation of Tension Wood Fibre in Fraxinus mandshurica Rupr. var. japonica Maxim.

The ultrastructure of the fibre wall in Fraxinus mandshuricaRupr. var. japonica Maxim. was investigated by electron microscopy.The trees had been inclined artificially at an angle of 30°to the vertical at the beginning of the initiation of cambialgrowth in early spring. The secondary walls of tension woodfibres were of the outer (S₁) layer and gelatinous (G) layertype. The microfibrils in the gelatinous (G) layer were orientedas a steep Z-helix relative to the fibre axis with a deviationthat ranged from 0° to 25° but was mainly between 5°and 10°. The cross-sectional surface of tension wood fibresrevealed the relatively strong attachment of the G-layer tothe S₁ layer. The G-layer stained weakly with potassium permanganate.The S₁ layer of tension wood fibres stained less strongly thanthat of the normal and opposite wood fibres. These results indicatethat the tension wood in F. mandshurica var. japonica is nottypical and is somewhat anomalous. The secondary walls of normaland opposite wood fibres were composed of two layers, S₁ andS₂, and lacked an S₃ layer. Microfibrils in the S₃ layer ofjuvenile stems were extremely variable in orientation and weresparsely distributed without forming a layer. By contrast, avery thin S₃ layer was present in the wood fibres of maturestems. The variations in the formation of the S₃ layer in thefibre walls were probably due to the differences in the cambialage of the stems of F. mandshurica Rupr. var. japonica.Copyright1995, 1999 Academic Press Fraxinus mandshurica Rupr. var. japonica Maxim., Japanese ash, tension wood, fibre wall, G-layer, microfibrillar orientation, normal and opposite wood, juvenile stem, field-emission scanning electron microscopy, low accelerating voltage     

自然条件下风箱果的克隆构型

在自然条件下,风箱果(Physocarpus amurensis)主要靠克隆繁殖维持种群。植物克隆构型的可塑性变化对于其适应环境异质性具有重要意义。为探求风箱果的克隆构型及根茎生长动态,研究了其地下根茎的构筑型、形态特征、根茎的直径随长度变化的规律和地下根茎间的夹角。结果表明:风箱果的地下根茎的构筑型基本上属于游击型;风箱果无性系平均含有(6±2)个分株和(9.33±3.48)个根茎;根茎的直径随长度变化的曲线为抛物线型;分枝夹角较为稳定,多为30°和70°。风箱果生产大量的根茎系统,每一个克隆片段能够占据一定的空间,以保证自身生存和维持种群繁衍。     

An Experimental Investigation of the Resistance of Model Root Systems to Uprooting

The architecture of a tree root system may influence its abilityto withstand uprooting by wind loading. To determine how theroot branching pattern may alter the anchorage efficiency ofa tree, artificial model root systems with different topologiesand branching angles were built. The root systems were embeddedat various depths in wet sand and the pull-out resistance measured.A model to predict the uprooting resistance from the data collectedwas designed, allowing predictions of anchorage strength withregards to architecture. The dominant factors influencing pull-outresistance were the depth and length of roots in the soil. Themost efficient type of branching pattern predicted by the programwas one with an increased number of roots deep in the soil.The optimum branching angle most likely to resist pull-out isa vertical angle of 90° between a lateral and the main axis.The predicted mechanically optimal radial angle between a lateralbranch and its daughter is between 0 and 20°. Values ofbranching angle are compared with those measured in real woodyroot systems of European larch and Sitka spruce. Root architecture; root anchorage; pull-out resistance; windthrow; Picea sitchensis ; Larix decidua     

Impacts of the invasive plant <Emphasis Type="Italic">Fallopia japonica</Emphasis> (Houtt.) on plant communities and ecosystem processes

Fallopia japonica (Japanese knotweed) invades riparian areas and roadsides in New England. This large clonal species drastically alters the appearance of habitats by forming highly productive near-monocultures. To understand how these invasions affect ecosystem processes in New England, we quantified the impacts of F. japonica on species diversity, primary productivity, and nitrogen cycling at five locations in central Massachusetts, USA. In stands of F. japonica and in adjacent uninvaded areas, we recorded the cover of each plant species and measured the aboveground biomass and nitrogen (N) concentrations in plants, along with N retranslocation from F. japonica leaves and several soil characteristics. In addition, we severed rhizomes of peripheral F. japonica shoots to determine if clonal integration contributes to the species’ rapid spread and dominance. Stands of F. japonica had lower species diversity, but greater aboveground biomass and standing N than uninvaded areas. Nitrogen and carbon concentrations in biomass and N mineralization rates in soil did not differ between stands and adjacent areas. Rhizome severing temporarily reduced growth of F. japonica, suggesting that retranslocation of photoassimilates and/or nutrients between shoots via rhizomatal connections may maximize stand level growth rates and facilitate dominance by F. japonica.     

Effects of Photoperiod and Temperature on Shoot, Root and Rhizome Growth in ThreeLotus uliginosusSchkuhr Populations

The growth of three populations of greater lotus (Lotus uliginosusSchkuhrsyn.L. pedunculatusCav.) was compared at photoperiods of 10,12 and 14 h at a maximum day/minimum night temperature of 21/16°C and at maximum day/minimum night temperatures of 27/22,21/16, 18/13 and 15/10 °C at a photoperiod of 12 h. Shortdays (10 h) favoured root and rhizome development compared tolong days (14 h). A temperature regime of 15/10 °C restrictedrhizome development compared to the 18/13 and 21/16 °C regimes.Shoot growth was restricted at the highest temperature regime(27/22 °C). The cultivar Sharnae had fewer, but heavier,rhizomes than Grasslands Maku; this may indicate adaptationto the dry summers at its site of origin (Algarve, Portugal).The response of rhizome growth to temperature and photoperiodexplains part of the performance of greater lotus in the fieldat a wide range of latitudes. Grazing management to encouragethe persistence ofL. uliginosusin pasture in temperate environmentsmay include the exclusion of grazing livestock in autumn. Inthe sub-tropics, monitoring of rhizome production in the fieldwould be required before deciding the appropriate time intervalbetween grazing.Copyright 1998 Annals of Botany Company Lotus uliginosus(Schkuhr); greater lotus; temperature; daylength; shoots; roots; rhizomes.     

Respiration for maintenance and growth inReynoutria japonica ecotypes from different altitudes on Mt Fuji

Respiration for maintenance and growth ofReynoutria japonica ecotypes from altitudes of 700 and 2420 m on Mt Fuji were measured in two controlled thermal conditions. The maintenance respiration of the high-altitude ecotype at both 15 and 25°C was significantly (1.7-fold) higher than that of the low-altitude ecotype, whereas growth respiration was independent of both ecotype and temperature. The temperature coefficient (Q₁₀) of the maintenance respiration was about 1.9 in both ecotypes. The results show that there is ecotypic differentiation in the performance of maintenance respiration. It is suggested that the high maintenance respiration of the high-altitudeR. japonica ecotype is advantageous in severe upland environments but disadvantageous in a warm lowland climate in terms of carbon economy.     

Ontogenetic changes in accumulation of rhizomes in monoclonal patch of <Emphasis Type="Italic">Miscanthus sinensis</Emphasis> Anderss. in warm temperate region of Japan

To determine the main benefits of clonal expansion of Miscanthus sinensis patches (monoclones), we observed the annual pattern of the areal expansion of a number of M. sinensis patches and examined how the quantity of rhizomes in such patches is related to changes in their basal area. To forage for nutriments, a patch must continuously widen its habitat. Patches annually expanded centrifugally by sympodial branching of short rhizomes, which originated in tillering that occurred more than once a year. However, the basal area of the patches approached a ceiling as the patches aged. Both the number and the weight of rhizomes in the patches continued to increase as long as the basal area expanded. The mean weight of rhizomes in patches also initially increased quickly, but then reached a ceiling as the clones expanded. Similarly, the amount of reserve substance per shoot in the patches increased asymptotically along with the clonal expansion, depending on the rhizome mass allotted to each shoot. These results suggest that, in the clonal growth of M. sinensis patches, the accumulation of reserve matter in the rhizomes is more important than foraging in new areas.     

Plant Growth and Survival under Strict Anaerobiosis

Anaerobic incubation of seedlings and rhizomes reveals interspecificdifferences in the ability of seedlings and rhizomes of higherplants to survive under prolonged and strict anaerobiosis. Rhizomesof several species were killed by an anaerobic incubation at22 °C for 7 d while others survived and showed normal shootextension on return to aerobic conditions. A third group ofspecies showed healthy and geotropically normal bud and shootextension while their rhizomes were in the oxygen free environment.A detailed comparison of bud and shoot growth rates was madewith Scirpus maritimus under aerobic and anaerobic conditionsover a 14 d period. Under similar conditions four species of grass seedlings weresubjected to anaerobic conditions for 2–8 d at 5°C,15 °C and 25 °C. Seedling mortality was highest at 25°C in all four grasses. Interspecific differences were evident.The wetland species were more tolerant of anoxia than the grassesfrom drier habitats.     

Effect of Light Quality (Red:Far-red Ratio) at the Apical Bud of the Main Stolon on Morphogenesis of Trifolium repens L.

A controlled environment experiment investigated whether thered:far-red (R:FR) ratio of light at the apical bud of the mainstolon could alter plant morphogenesis in clonal cuttings ofwhite clover (Trifolium repens L.) The apical bud included theapical meristem, five to six developing leaf primordia withassociated axillary bud primordia and stipules and the firstemerged folded leaf until development was greater than 0·3on the Carlson scale. Three light regimes were imposed on theapical bud by collimating light from R or FR light-emittingdiodes so that the R:FR ratio of light incident at the apicalbud was set at 0·25, 1·6 or 2·1, withoutsignificantly altering photosynthetically active radiation.The effect of these light regimes on white clover seedling growthwas also tested. At a low R:FR ratio seedling hypocotyl and cotyledon lengthswere significantly longer. However, with the cuttings, the lighttreatments did not alter node appearance rate or internode lengthof the main stolon, petiole length, area of leaves or totalshoot dry matter. There was one significant photomorphogeneticresponse in the cuttings, a delay of 0·5 of a phyllochronin the appearance of branches from axillary buds in the lowR:FR ratio treatment relative to the other treatments. Wherebranch appearance was delayed plants had fewer branches. Thisdifference could be ascribed solely to a delay in branch appearanceas there were no significant treatment effects on either theinitiation of axillary bud primordia within the apical bud,the probability of branching or on the rate of growth of branchesafter appearance. Because treatment of the apical bud inducedonly one of the many previously observed responses of whiteclover to a decrease in the R:FR ratio of light, we concludethat other plant organs must also sense the quality of incidentlight.Copyright 1994, 1999 Academic Press White clover, Trifolium repens, apical bud, light quality, red:far-red ratio, light-emitting diode, branching, axillary buds, photomorphogenesis     

Gravitropism of Primary Roots of Zea mays L. at Different Displacement Angles

Primary roots of Zea mays were oriented at various angles fromthe vertical ranging from 99° to 1° and their subsequentbending analysed from filmed records. The maximum rate of bendingand the time before bending commenced both varied two-fold,but showed no correlation with the initial angle of tip displacement.Roots orientated to small initial angles (< 40°) oftenovershot the vertical and proceeded to oscillate around thisorientation, whereas roots oriented to large initial angles(> 60°) often failed to achieve the vertical. Roots inthis latter group resumed bending after an indeterminate time,or did so immediately after a second displacement of their tip,showing that they were not intrinsically unable to bend. Theapparently spontaneous resumption of bending after a temporaryplagiogravitropic phase is suggested as being due to noise inthe graviperception system in the root cap. The tips of rootsgrowing vertically downwards showed oscillatory bending movementsup to 10° either side of vertical. This angle correspondsto the minimum angle of displacement which induces gravitropicbending. Only when roots were oriented 10-20° from verticaldid they begin unequivocally to show a gravitropism since atsuch angles the deflection of their tips exceeded that due totheir natural oscillation.Copyright 1993, 1999 Academic Press Gravitropism, roots, Zea mays     

Sporulation in Rhizopus sexualis and Some Other Fungi Following a Period of Intense Respiration

Rhizopus sexualis grown at 20° C. on liquid I per cent.malt or glucose-asparagine medium showed a peak of respiratoryactivity between 40 and 55 hours-after inoculation. Rate ofrespiration then fell until it reached a steady low level whichcoincided with maximum mycelial growth. Zygospore initiationoccurred at or just after the peak of respiration. At a low temperature (9° C.) or with high concentrationsof glucose the respiration peak was less marked and no zygosporesdeveloped. Single ‘plus’ or ‘minus’strains of the heterothallic species Mucor hiemalis and Phycomycesblakesleeanus showed a pattern of respiration and mycelial growthsimilar to that of R. sexualis but no zygospores were formed.Zygospores did not develop without a preliminary period of intenserespiration, but such a peak period could occur without beingfollowed by zygospore formation. A strain of Sordaria fimicola was grown at 25° C. on a syntheticmedium with 5.0 per cent. sucrose or glucose as source of carbon.Respiration reached a peak at approximately 4 and 5 days respectively,the actual peak value being twice as large on sucrose as onglucose. Dry weight of mycelium was greater on glucose thanon sucrose. Perithecia were formed only on the sucrose medium.Visible peri-thecial initials were first seen shortly afterthe occurrence of the respiratory peak. Mature perithecia werepresent 3 days later. The possible significance of these results is discussed.     

Effects of Temperature and Photoperiod on Growth and Development of Tillers and Rhizomes in Poa pratensis L. Ecotypes

The vegetative growth of four Norwegian ecotypes of Poa pratensisL. was compared at day/night temperatures of 21/12, 21/6, 12/12and 12/6 °C and at photoperiods of 12, 16, 20 and 24 h,the irradiation being approximately equal in all treatments.Tillering within tufts was most abundant in short days and atthe large temperature amplitude. Rhizome formation and elongationwere stimulated by long days and, more strongly, by high daytemperature, but a greater proportion of the rhizomes formedaerial tillers in short days. Long day stimulation of heightgrowth and dry matter accumulation differed between ecotypesbut was generally most pronounced at low temperatures. The NorthNorwegian ecotype ‘Lavang’ had a higher relativegrowth rate and developed two to three times as many rhizomesas its South Norwegian counterparts. Day/night temperature, growth rate, leaf area, photoperiodicity, Poa pratensis L., rhizomes, roots, smooth meadow grass, tillering, weight gain     

The Hexagonal Branching Pattern of Rhizomes of Alpinia speciosa L. (Zingiberaceae)

Alpinia speciosa L. (Zingiberaceae) is a striking example ofa plant in which an organized architectural structure fulfilsa basic functional requirement - the economical explorationand exploitation of the substrate. This rhizomatous plant spreadslaterally in a predictable manner by vegetative growth to forma clone of separated ramets. A morphological study establishes‘rules of growth’ governing the success and failureof lateral branches (sympodial units), their lengths, and theirrelative angles. The basic branching pattern of this rhizome system approximatesto that of a hexagonal grid - the most economical structuralshape that uniformly ‘samples’ a plane surface.It is shown that the significant deviation of branching anglefound in Alpinia from that required to develop an exact hexagonalnetwork will virtually eliminate aerial shoot ‘clashes’in which the placing of one aerial shoot coincides with thatof another aerial shoot, whilst at the same time enhancing thebenefits of the hexagonal system. The broader implications ofthe organised architecture of plants are emphasised. rhizome, branching pattern, architecture, productivity     

The Galls on Forsythia intermedia Zab.

The galls on Forsythia intermedia Zab. consist of much-branched,root-like structures embedded in corky material which appearsto be sloughed-off from them. These gall ‘roots’have 7 to 14 vascular bundles compared with the 5 of normalroots and contain much more IAA. When galls were incubated in damp chambers the gall ‘roots’extended their growth. These extensions had a root-cap and root-hairsbut also had more vascular bundles than normal and they didnot respond geotropically like normal roots. Two fungi, Gibberella baccata (Wallr.) Sacc. (conid, stat. Fusarlumlateritium Nees.) and Phomopsis dominici Trav. were associatedwith die-back of gall ‘roots’ and two bacteria resemblingCorynebacterium fascians (Til.) Dowson and Agrobacterium tumefaciens(Sm. and Towns.) Conn respectively were isolated from galls.All produced IAA in culture media but their role in the etiologyof the gall, if any, remains in doubt.     

Cold Acclimation, Freezing Resistance and Protein Synthesis in Alfalfa (Medicago sativaL. cv. Saranac)

Mohapatra, S. S., Poole, R. J. and Dhindsa, R. S. 1987. Coldacclimation, freezing resistance and protein synthesis in alfalfa(Medicago sativa L. cv. Saranac).—J. exp. Bot. 38: 1697–1703. Changes in freezing resistance (percent survival at —10°C), pattern of protein synthesis and translatable mRNApopulation during cold acclimation of alfalfa (Medicago sativaL. cv. Saranac) have been examined. Two days of cold acclimationat 4 °C increased freezing resistance from about 6% to 40%,protein content by 200% and total RNA content by 100%. Acclimationfor longer periods did not cause further increases in freezingresistance, protein content or RNA content. Examination of proteinchanges by sodium dodecyl sulphate-polyacrylamide gel electrophoresis(SDS-PAGE) coupled with protein staining, and by fluorographyof in vivo labelled proteins separated by SDS-PAGE, showed thatseveral proteins are increasingly or newly synthesized duringcold acclimation. Analysis of in vitro translation productsby SDS-PAGE and fluorography shows changes in the populationof translatable mRNAs. It is concluded that in this varietyof alfalfa cold acclimation for only 2 d is sufficient to confermaximum freezing resistance, and that changes in proteins duringcold acclimation are regulated most probably at the transcnptionallevel. Key words: Freezing resistance, protein synthesis, cold acclimation, SDS-PAGE, Medicago sativa L.     

Seed Production Environment, Time of Harvest, and the Potential Longevity of Seeds of Three Cultivars of Rice (Oryza sativa L.)

Changes in seed quality (assessed by potential longevity, i.e.the value of the seed lot constant K₁ of the seed viabilityequation) in three contrasting cultivars of rice (Oryza sativaL.) were monitored during seed development and maturation intwo temperature regimes, viz 28/20°C and 32/24°C (12/12h), provided by controlled environments. Mass maturity (definedas the end of the seed-filling phase) varied only between 18and 20 d after 50% anthesis. In five of the six treatment combinationsmaximum potential longevity was not achieved until 12-19 d aftermass maturity. In contrast, the maximum potential longevityof seeds of a japonica rice cultivar produced in the warmerregime was obtained in the first harvest after mass maturity.After mass maturity, the potential longevity of the japonicarice seed lots produced in the warmer environment was much lessthan that for the cooler environments. Maximum potential longevitywas also consistently greater in the cooler than the warmerregime for the two indica cultivars, although the differencein K₁ was small (0·3-0·5). The deleterious effectof increase in temperature on seed quality development was notdetected until after mass maturity. Maximum potential longevityin the cooler regime was greatest in the glutinous indica (K₁= 3·9) and least in the japonica cultivar (K₁ = 3·1).It is concluded that the japonica cultivar is not as well adaptedto warm seed production regimes as the indica cultivars. Consequently,subject to confirmation, this research suggest that the seedproduction of japonica cultivars for long-term genetic conservationshould be undertaken, whenever possible, in warm temperate environments.Copyright1993, 1999 Academic Press Oryza sativa L., rice, genebanks, seed development, seed storage, seed longevity, temperature     

Relative costs and benefits of clonal integration of Zoysia japonica under various N:P ratios

Clonal plants adaptively respond to abiotic stress, but to date little is known about under what circumstanses clonal integration is beneficial, or costly. To study the costs and benefits of clonal integration on clonal growth, we cultivated Zoysia japonica under three ratios of N:P (N:P ≈ 7, 14:1 and 21:1), and four types of stolon severing treatments (connected stolon CS, light severing LS, moderate severing MS, serious severing SS). The results showed that Z. japonica performed best at a low ratio of N:P (N:P ≈ 7:1). When the stolons were connected (CS), the growth of primary A‐ramets, multiple‐nodes and stolons benefited from clonal integration; however, the growth of primary B‐ramets on the primary stolons, A and B ramets was significantly reduced (p < 0.05). In the moderate stolon severing treatment MS_7:1 (N:P ≈ 7:1), clonal integration appeared more cost effective than in all other treatments. On the whole, with increasing stolon severing intensity, the cost of clonal integration increased and the clonal growth of Z. japonica declined significantly. The pattern of biomass allocation may be useful for Z. japonica to adapt to the various environments, and clonal integration plays a significant role under adverse environmental conditions.