Growth-induced water potentials may mobilize internal water for growth

Abstract. Wphen there is no external source of water, plants can grow by mobilizing internal water from nongrowing tissues. We investigated how this internal water moves by measuring continuously and simultaneously the water potential (ψ_w) of soybean ( Glycine max L. Merr.) seedlings in the upper, growing stem tissues and the lower, non-growing stem tissues. When external water was available to the roots, the stems grew rapidly and the ψ_w of the growing tissue was continually below that of the nongrowing tissue and the medium around the roots. This indicated that a growth-induced gradient in ψ_w favoured water movement from the external source to the growing cells. When the external source was removed, the ψ_w of the growing tissue remained constant for a time and the ψ_w of the nongrowing tissue decreased somewhat. Growth took place slowly as water was withdrawn from the nongrowing tissue but ψ_w gradients continued to favour water transport to the growing cells. On the other hand, if this internal source was removed by excision, growth ceased abruptly. In this case, the cell walls relaxed and the ψ_w of the growing tissue decreased by about 0.1 MPa instead of remaining constant. The ψ_w of the detached nongrowing tissues remained constant instead of decreasing. This indicates not only that water mobilization required attached nongrowing or slowly growing tissues but also that mobilization affected wall relaxation. Thus, ψ_w differences may mobilize internal water, may explain the continued growth of plants and plant parts removed from external sources of water, and may account for discrepancies in measurements of cell wall properties in growing tissues.     

Osmoregulation,solute distribution,and growth in soybean seedlings having low water potentials

Soybean (Glycine max (L.) Merr.) seedlings osmoregulate when the supply of water is limited around the roots. The osmoregulation involves solute accumulation (osmotic adjustment) by the elongating region of the hypocotyls. We investigated the relationship between growth, solute accumulation, and the partitioning of solutes during osmoregulation. Darkgrown seedlings were transplanted to vermiculite containing 1/8 (0.13 x) the water of the controls. Within 12–15 h, the osmotic potential of the elongating region had decreased to-12 bar, but it was-7 bar in the controls. This osmoregulation involved a true solute accumulation by the hypocotyls, since cell volume and turgor were virtually the same regardless of the water regime. The hypocotyls having low water potentials elongated slowly but, when deprived of their cotyledons, did not elongate or accumulate solute. This result indicated a cotyledonary origin for the solutes and a dependence of slow growth on osmotic adjustment. The translocation of nonrespired dry matter from the cotyledons to the seedling axis was unaffected by the availability of water, but partitioning was altered. In the first 12 h, dry matter accumulated in the elongating region of the 0.13 x hypocotyls, and osmotic adjustment occurred. The solutes involved were mostly free amino acids, glucose, fructose, and sucrose, and these accounted for most of the increased dry weight. After osmotic adjustment was complete, dry matter ceased to accumulate in the hypocotyls and bypassed them to accumulate in the roots, which grew faster than the control roots. The proliferation of the roots resulted in an increased root/shoot ratio, a common response of plants to dry conditions.Osmotic adjustment occurred in the elongating region of the hypocotyls because solute utilization for growth decreased while solute uptake continued. Adjustment was completed when solute uptake subsequently decreased, and uptake then balanced utilization. The control of osmotic adjustment was therefore the rate of solute utilization and, secondarily, the rate of solute uptake. Elongation was inhibited by unknown factors(s) despite the turgor and substrates associated with osmotic adjustment. The remaining slow elongation depended on osmotic adjustment and represented some optimum between the necessary inhibition for solute accumulation and the necessary growth for seedling establishment.     

THE PHYSIOLOGICAL BASIS OF MORPHOLOGICAL POLARITY IN REGENERATION. II

1. The experiments show that the mass of air roots formed in a stem increases with the mass of the leaf attached to the stem, though it has not been possible to establish an exact mathematical relation between the two masses, owing to unavoidable sources of error. 2. Darkened leaves do not increase the mass of roots formed. 3. In stems suspended horizontally air roots appear on the lower side of the stem, with the exception of the cut end where they usually appear around the whole circumference of the stem. When the lower half of a stem suspended horizontally is cut off, roots are formed on the upper side. It is shown by experiments on leaves suspended horizontally that the more rapidly growing roots and shoots on the lower side inhibit the root and shoot formation in the upper half of such a leaf; and likewise the more rapid formation of roots on the lower side of a horizontally suspended stem seems to account for the inhibition of root formation on the upper side of such a stem. Likewise the more rapid growth of shoots on the upper side of a stem suspended horizontally is likely to inhibit the growth of shoots on the lower side. 4. Each leaf contains in its axil a preformed bud capable of giving rise to a root, which never grows out in the normal stem on account of the inhibitory influence of the normal roots at the base of the plant. These dormant root buds are situated above (apically from) the dormant shoot bud. The apical root buds can be caused to develop into air roots when a piece of stem is cut out from a plant from which the leaves except those in the basal node of the piece are removed. The larger these basal leaves the better the experiments succeed. 5. These apical air roots grow out in a few days, while the roots at the basal end of the stem (which in our experiments dip into water) grow out about a week later. As soon as the basal roots grow out in water they cause the air roots in the more apical region of the stem to dry out and to disappear. 6. In addition to the basal roots, basal nodes have also an inhibitory effect on the growth of the dormant root buds in the apical region of a stem. This is indicated by the fact that a stem with one pair of leaves near the base will form apical air roots more readily when no node is situated on the stem basally from the leaf than if there is a node basally from the leaf.     

Rapid,localized light effect on root growth in maize

A method using optical microfibers permitted localized exposure of the cap or the elongating part of growing maize (Zea mays L.) roots to white light. When the cap was illuminated, a strong and very rapid inhibition of the elongation rate of the roots was found. When the light microbeam was directed at the elongating region, the roots continued to grow at the same rate as before the illumination.     

Polysomes, Messenger RNA, and Growth in Soybean Stems during Development and Water Deficit

The polysome status and populations of polysomal mRNA were examined in different regions of dark-grown soybean (Glycine max [L.] Merr.) stems that contained either dividing, elongating, or mature (nongrowing) cells. There was a developmental gradient of polysome content in which the dividing tissue had the highest levels and the mature tissue the lowest. A few hours after transplanting the seedlings to vermiculite having low water content (water potential Ψ_w = −0.29 megapascals), stem growth rate decreased to 30% of well-watered controls and the polysome content decreased most in the dividing and elongating tissues. After 24 to 36 hours, stem growth and polysome content recovered gradually. In vitro translation products of polysomal mRNA from dividing, elongating or mature tissue were examined on two-dimensional gels. In well-watered controls, each of the stem regions was enriched in a small subset of the polysomal mRNA population, probably because of developmentally regulated gene expression. Exposing plants to low Ψ_w for 24 hours induced a change in the relative abundance of a small number of polysomal mRNAs in the elongating and mature tissues, but not in the dividing tissue. After 24 to 72 hours at low Ψ_w, the changes in polysomal mRNA population were reversed in the elongating tissue. The data indicate that changes in stem growth at low water potential are associated with changes in polysome status and polysomal mRNA in the elongating tissue.     

Turgor and growth at low water potentials

Turgor affects cell enlargement but has not been measured in enlarging tissue of intact plants when growth is inhibited by inadequate water. Mature or excised tissue can be problematic for these measurements because turgor may not be the same as in intact enlarging cells. Therefore, we measured the average turgor in the elongating region of intact stems of soybean (Glycine max [L.] Merr.) while the seedlings were exposed to low water potentials by transplanting to vermiculite of low water content. Stem growth was completely inhibited by the transplanting, and the average turgor decreased in the mature stem tissue. However, it did not decrease in the elongating region whether measured in intact or excised tissue (total of four methods). At the cellular level, turgor was uniform in the elongating tissue except at transplanting, when turgor decreased in a small number of cortical cells near the xylem. The reduced turgor in these cells, but constant turgor in most of the cells, confirmed that no general turgor loss had occurred but indicated that gradients in water potential extending from the xylem into the enlarging tissue were reduced, thus decreasing the movement of water into the tissue for cell enlargement. A modest growth recovery occurred after 2 days and was preceded by a recovery of the gradient. This suggests that under these conditions, growth initially was inhibited not by turgor loss but by a collapse of the water potential gradient necessary for the growth process.     

Cell wall proteins at low water potentials

We investigated the proteins extractable from cell walls of stem tissues when plants were subjected to low water potentials (low ψ_w). Dark-grown soybean seedlings (Glycine max [L.] Merr.) showed decreased stem growth when the roots were exposed to vermiculite having low water content (ψ_w = −3 bar). After a time, growth resumed but at a reduced rate relative to the controls. The extractable protein increased in the cell walls as ψ_w decreased, especially a 28-kilodalton protein in the young tissue. In contrast, a 70 kilodalton protein, mainly extractable from mature cell walls, appeared to decrease slightly at low ψ_w. No hydroxyproline was present in either protein, which shows that neither protein is related to extensin. The level of the 28 kilodalton protein increased in the cell wall of the dividing region soon after the initial growth inhibition, and it appeared in the elongating tissue at about the time growth resumed. The correlation between growth and these protein changes suggests that the two events could be related.     

Direct Demonstration of a Growth-Induced Water Potential Gradient

When transpiration is negligible, water potentials in growing tissues are less than those in mature tissues and have been predicted to form gradients that move water into the enlarging cells. To determine directly whether the gradients exist, we measured water potentials along the radius of stems of intact soybean (Glycine max [L.] Merr.) seedlings growing in vermiculite in a water-saturated atmosphere. The measurements were made in individual cells by first determining the turgor with a miniature pressure probe, then determining the osmotic potential of solution from the same cell, and finally summing the two potentials. The osmotic potentials were corrected for sample mixing in the probe. The measurements were checked with a thermocouple psychrometer that gave average tissue water potentials. In the elongating region, the water potential was highest near the xylem and lowest near the epidermis and in the center of the pith. In the basal, more mature region of the same stems, water potentials were near zero next to the xylem and throughout the tissue. These basal potentials reflected mostly the potential of the xylem, which extended into the elongating tissues. Thus, the high basal potential confirmed the high potential near the xylem in the elongating tissues. The psychrometer measurements for each tissue gave average potentials that agreed with the average of the cell potentials from the pressure probe. We conclude that a radial gradient was present in the elongating region that formed a water potential field in three dimensions around the xylem and that confirmed the predictions of Molz and Boyer (F.J. Molz and J.S. Boyer [1978] Plant Physiol 62: 423-429).     

Low water potentials affect expression of genes encoding vegetative storage proteins and plasma membrane proton ATPase in soybean

We have examined growth, water status and gene expression in dark-grown soybean (Glycine max L. Merr.) seedlings in response to water deficit (low water potentials) during the first days following germination. The genes encoded the plasma membrane proton ATPase and two proteins of 28 kDa and 31 kDa putatively involved in vegetative storage. Water potentials of stems and roots decreased when 2-day-old seedlings were transferred to water-saturated air. Stem growth was inhibited immediately. Root growth continued at control rates for one day and then was totally inhibited when the normal root-stem water potential gradient was reversed.Expression of mRNA for the 28 kDa and 31 kDa proteins, measured independently using specific 3-end probes, occurred about equally in stems. However, only the mRNA for the 31 kDa protein was detected in roots and at a lower abundance than in stems. Low water potentials increased the mRNA only for the 28 kDa protein in stems and the 31 kDa protein in roots. This differential expression followed the inhibition of stem growth but preceded the inhibition of root growth.The expression of the message for the ATPase, measured using a probe synthesized from a partial oat ATPase clone, was low in stems and roots but there was a 6-fold increase at low water potentials in roots. The increase followed the inhibition of root growth. This appears to be the first instance of regulation of ATPase gene expression in plants and the first demonstration of differential expression of the 28 kDa, 31 kDa, and ATPase messages. The correlation with the differential growth responses of the stems and roots raises the possibility that the differential gene expression could be involved in the growth response to low water potentials.     

土壤湿度对香樟幼苗根系生长和构型的影响

以1年生香樟(Cinnamomum camphora)幼苗为试材,设置对照组(CK)、中度干旱处理(M)、重度干旱处理(S)三个处理,比较不同土壤湿度下香樟幼苗不同时期地上部分生长和根系构型,探究香樟幼苗根系对不同土壤湿度的适应性及其耐旱机制。结果表明,中度和重度干旱处理组的香樟根系及地上部分干物质积累、根系长度、根系表面积、根系直径和根尖数均显著小于对照组(P<0.05)。同时干旱显著增加香樟幼苗的根系拓扑指数,降低香樟根的分形维数和平均分枝角度(P<0.05)。可见土壤湿度程度及处理时间显著影响香樟根系的生长及在土壤中的布局。较低土壤湿度可显著抑制根长的延长、根表面积扩大和根的增殖,且随着土壤湿度的继续降低以及处理时间延长,香樟幼苗根系的生长受到水分亏缺的抑制作用加重,根系建成成本增高的同时,根系分枝的复杂性降低,根系必须通过朝着更陡、更深的方向生长伸长来提高水分吸收效率。建议在园林绿化工程养护过程中制定科学的水分管理策略,以满足香樟生长过程中对土壤水分的需要。     

The root tip and accelerating region suppress elongation of the decelerating region without any effects on cell turgor in primary roots of maize under water stress

To identify the region in which a root perceives a decrease in the ambient water potential and changes its elongation rate, we applied two agar blocks (1 x 1 x 1 mm(3)) with low water potential bilaterally to primary roots of maize (Zea mays) at various positions along the root. When agar blocks with a water potential of -1.60 MPa (-1.60-MPa blocks) or lower were attached to a root tip, the rate of elongation decreased. This decrease did not result from any changes in the water status of elongating cells and was not reversed when the -1.60-MPa blocks were replaced by -0.03-MPa blocks. The rate decreased slightly and was unaffected, respectively, when -1.60-MPa blocks were applied to the so-called decelerating region of the elongating zone and the mature region. However, the rate decreased markedly and did not recover for several hours at least when such blocks were attached to the accelerating region. In this case, the turgor pressure of the elongating cells decreased immediately after the application of the blocks and recovered thereafter. The decrease in elongation rate caused by -1.60-MPa blocks applied to the root tip was unaffected by additional -0.03-MPa blocks applied to the accelerating region and vice versa. We concluded that a significant reduction in root growth could be induced by water stress at the root tip, as well as in the accelerating region of the elongating zone, and that transmission of some signal from these regions to the decelerating region might contribute to the suppression of cell elongation in the elongation region.     

Laser ablation ICP-MS reveals patterns of copper differing from zinc in growth zones of cucumber roots

Laser ablation coupled with inductively coupled plasma-mass spectrometry was used to find Cu and Zn concentration in surface tissue along a longitudinal developmental gradient with meristem, rapidly elongating tissue, and nongrowing tissue in a model system of seedling roots of Cucumis sativus L. (cucumber). Tissue metal accumulation was determined for roots of seedlings growing on cellulosic germination paper treated with nutrient solution (controls), and also treated with concentrations of Zn (40 ppm) and Cu (10 ppm) that reduced growth. Cu content of all roots is highest at the apex and falls sharply to lower values by 2 mm from the root tip. In contrast, at moderate Zn availability (0.07 ppm), Zn content rises from the apex to 2 mm then falls throughout the remainder of the growth zone. At high external Zn the spatial pattern resembles that of Cu. Cucumber root growth zones accumulate more of each metal with higher external availability. Metal deposition rates were calculated using a continuity equation with data on local metal content and growth velocity. Deposition rates of both metals are generally highest in the rapidly elongating region, 1.5–3.5 mm, even where metal concentration is decreasing with position and root age and even when the accumulation is inhibitory to growth.     

Interpreting changes in stem diameter in rose plants

Sixteen adult rose plants ( Rosa hybrida cv. Sweet Promise) were grown in rockwool in a phytotron. Water uptake was estimated by computing the difference between the rate of nutrient solution supply and the rate of leaching, continuously measured using a high-precision flowmeter. Transpiration was measured using a balance continuously recording the loss of weight in the entire system. Changes in stem diameter were recorded using linear variable differential transducers placed at the base of four developing shoots and on their corresponding basal stems. Changes in stem diameter were related to changes in the water content of 16 plants. The amplitude of shrinkage in daytime stem diameter and daily increase in stem diameter was lower in basal stems. Radial hydraulic resistance to water diffusion in stems seems to be very low and stem reservoirs appear to be very early sensors of water depletion. When the nutrient solution supply was suppressed, the amplitude of the daily increase in stem diameter and transpiration rates were reduced one and one-and-a-half days later, respectively. When water depletion became more severe, stem diameter decreased from one day to another. The amplitude of daily stem shrinkage increased sharply for 6 days after watering was stopped, while the bulk modulus of elasticity of leaves decreased, probably as a result of damage to the cell wall.     

The Casparian strip in pea epicotyls: Effects of light on its development

The Casparian strip, which is specific to roots, was studied in the epicotyls of dark-grown seedlings of pea (Pisum sativum L.) where it was found to have the same morphology and properties as the strip in roots. In dark-grown seedlings, the distance between the upper-most position of the Casparian strip and the bending point of the hook (about 37 mm) did not change during growth of the seedlings. In the uppermost 0.5-mm region of the region in which the Casparian strip could be detected by fluorescence microscopy, the plasma membrane was not firmly attached to the cell wall. The development of the Casparian strip continued for about 42 h after dark-grown seedlings were transferred to the light, indicating that (i) the cells that have been determined to form the Casparian strip in darkness form the strip in the light, and that (ii) it takes about 42 h for the cells to complete formation of the strip. Cells in the hook of dark-grown seedlings did not form a Casparian strip when such seedlings were transferred to the light. The Casparian strip was formed in rapidly elongating internodes of light-grown seedlings when the seedlings were transferred to darkness. Light did not control the formation of the Casparian strip in roots.Abbreviation PBS phosphate-buffered saline     

Water potentials induced by growth in soybean hypocotyls

Gradients in water potential form the driving force for the movement of water for cell enlargement. In stems, they are oriented radially around the vascular system but should also be present along the stem. To test this possibility, growth, water potential, osmotic potential, and turgor were determined at intervals along the length of dark-grown soybean (Glycine max L. Merr., cv. Wayne) hypocotyls. Transpiration was negligible in the dark, humid conditions, so that all water uptake was for growth. Elongation occurred in the terminal 1.5 centimeters of the hypocotyl. Water potential was −3.5 bars in the elongating region but −0.5 bar in the mature region, both in intact plants and detached tissue. There was a gradual transition between these values that was related to the growth profile along the hypocotyl. Tissue osmotic potentials generally paralleled tissue water potentials, so that turgor was the same throughout the length of the hypocotyl. If the elongating zone was excised, growth ceased immediately. If the elongating zone was excised along with mature tissue, however, growth continued, which confirmed the presence of a water-potential gradient that caused longitudinal water movement from the mature zone to the elongating zone. When the plants were grown in vermiculite having low water potentials, tissue water potentials and osmotic potentials both decreased, so that water potential gradients and turgor remained undiminished. It is concluded that growth-induced water potentials reflect the local activity for cell enlargement and are supported by appropriate osmotic potentials.     

短期增温对贡嘎山峨眉冷杉幼苗生长及其CNP化学计量学特征的影响

2009年5月至10月,在中国科学院贡嘎山高山森林生态系统观测站附近,采用红外灯加热人工模拟气候变暖研究了增温对峨眉冷杉(Abies fabiri (Mast) Craib)幼苗生长和养分及其化学计量特征的影响。由于红外灯的增温作用,在幼苗的整个生长季节,增温样地地表下5 cm、10 cm、20 cm处土温平均高于对照样地5.04 ℃、4.81 ℃、4.35 ℃,而土壤含水量则分别降低了7.03%、6.10%、6.40%;地表20 cm处空气温度相比对照样地上升了1.12 ℃,而空气相对湿度则降低了6.30%。除茎重比外,增温处理降低了峨眉冷杉幼苗的根长、基径、株高、总生物量、根重比、叶重比、根冠比和比叶面积。经方差分析发现,增温处理后幼苗根、茎和叶的C平均含量与对照差异性均不显著(P>0.05),其中除茎的提高了2.76%外,根和叶分别降低了7.15%和2.29%;N平均含量除茎显著降低之外(P<0.05),根、叶分别提高了9.78%和5.70%;幼苗根、茎、叶的P平均含量均低于对照11.97%、10.69%和2.99%,并且根和茎与对照存在显著性差异(P<0.05)。增温处理后幼苗根、茎、叶各器官的C︰N、C︰P 、N︰P与对照均不存在显著性差异(P>0.05),其中C︰P均大于对照,而C︰N和N︰P与对照相比,均有不同程度的减小;C︰N、N︰P和C︰P的平均值(标准差)大小顺序依次为茎(92.594.92)>根(61.891.65)>叶(60.813.23)、叶(4.990.22)>根(4.440.58)>茎(3.640.10)和茎(336.358.70)>叶(302.854.49)>根(274.865.27)。实验结果表明：增温对幼苗生长和生物量积累具有明显的限制作用,对叶片生长的阻碍作用尤为突出;增温改变了幼苗根茎叶的CNP含量及其化学计量比格局;在养分供应上,增温和对照处理下幼苗生长均受N素限制。     

Water relations of rough lemon (Citrus jambhiri Lush.) citrus seedlings infected withFusarium solani

Summary Rough lemon citrus seedlings were inoculated withFusarium solani and evaluated for changes in water relations of leaves, stems, and roots. Inoculated seedlings had decreased leaf stomatal conductance, lower leaf water potential, lower water content, and higher leaf osmotic values compared to healthy plants. Visible wilt symptoms occurred as early as 24 h after inoculation. Transpiration and root conductivity were lower in diseased plants but stem conductivity in diseased plants did not differ from the control. Thus, wilting appears to be due to the inability of roots to supply water to the leaves.     

沙地樟子松幼苗干旱致死过程中非结构性碳水化合物的变化

以2年生沙地樟子松幼苗为对象,通过持续自然干旱处理,研究当土壤含水量下降到田间持水量的60%、40%、30%、20%和15%时幼苗叶片水势及不同器官(一年生叶、当年生叶、茎、粗根和细根)的可溶性糖、淀粉和非结构性碳水化合物(NSC)的含量,分析沙地樟子松幼苗在干旱致死过程中各器官NSC的分配规律及其适应机制.结果表明: 土壤含水量从田间持水量的40%下降到15%,幼苗叶片凌晨及正午水势无显著变化.当土壤含水量从田间持水量的60%下降到30%,各器官可溶性糖、淀粉、NSC含量和可溶性糖/淀粉先下降后上升.从30%下降到20%,当年生叶、一年生叶、茎和细根可溶性糖、淀粉和NSC含量降低,而粗根可溶性糖含量增加,淀粉和NSC含量减少.从20%下降到15%,当年生叶、一年生叶和茎可溶性糖、淀粉和NSC含量降低,粗根可溶性糖和NSC含量下降,淀粉含量上升,细根可溶性糖含量减少,淀粉和NSC含量增加.沙地樟子松幼苗通过不断调整各器官NSC及其组分含量变化以适应不同干旱环境,土壤含水量下降到田间持水量的30%后,幼苗可溶性糖和NSC含量总体呈下降趋势,淀粉在粗根和细根中积累,幼苗可能因碳耗竭而死亡.     

Growth-induced water potentials originate from wall yielding during growth.

Multicellular plants display growth-induced water potentials that generate tensions on water in the apoplast and move water into the growing cells. The potentials are sometimes assumed to arise from wall yielding, keeping the turgor pressure below what otherwise would occur. There has been no direct test of this theory, and therefore whole plants or growing regions of stems (hypocotyls) of dark-grown soybean (Glycine max L. Merr.) seedlings were sealed in a pressure chamber, and wall yielding was decreased by applying external pressure. In whole plants, external pressure had little effect because the plants and water supply were uniformly exposed to the pressure. If pressure was applied to the stem while the roots were outside in water, stem elongation was markedly inhibited because the pressure raised the water potential of the growing region and decreased water entry, reducing wall yielding. Further increasing the pressure prevented water entry completely and measured the tensions in the apoplast in the same growing regions. Tensions were about 0.19 MPa at low external pressure, but diminished as wall yielding was inhibited. At external pressures of about 0.63 MPa, wall yielding was abolished and tensions approached zero. There was a linear relation between wall yielding and tension, supporting the theory that wall yielding lowers the turgor thus causing most of the growth-induced water potential.     

水分因素对沙地柏实生苗水分和生长特征的影响

为探讨未来降雨变化对半干旱气候区毛乌素沙地常绿优势灌木沙地柏(Sabina vulgaris)实生苗水分和生长特征的可能影响，在鄂尔多斯沙地草地生态站开展了模拟降雨变化的实验。水分饱和亏缺和组织密度随土壤水分含量提高而降低，失水系数却相反，这表明沙地柏实生苗的保水抗旱性随模拟降雨量增加而降低。水分梯度对枝茎面积比无显著影响，叶质量茎面积比和叶质量枝面积比随土壤水分含量增大而降低，这意味着实生苗分枝的供水潜力随水分可利用性提高而降低。水分变化显著影响生物量分配，而对形态和植冠生产力指数均无显著影响；生物量、株高和基茎增量的变化反映了沙地柏生长的缓慢性。根系对水分变化的敏感性高于叶和茎，地下部分生物量投资随水分可利用性提高而增大，这暗示沙地柏实生苗可能通过节约利用水分方式适应自然生境中的水分胁迫。