Early growth and ecophysiological responses of Koa (Acacia koa A. Gray) seedlings to reduced water and phosphorus

Sites in need of restoration typically have one or more environmental factors that limit seedling establishment. Identifying ecophysiological responses to environmental stressors can provide important insights into mitigating measures that would allow seedlings to overcome such constraints to survival. Koa (Acacia koa A. Gray) is a nitrogen-fixing tree species endemic to Hawai?i that is highly valued in restoring degraded forest ecosystems, which are often limited in available water and phosphorus. This study examined how koa seedlings respond to conditions of reduced water (65 W) and no phosphorus (0P). After 17 weeks, seedlings subjected to 65 W or 0P accumulated less biomass, smaller root-collar diameters, and lower nitrogen and phosphorus contents. Combined reductions in water and P resulted in seedlings with increased root to shoot dry biomass and shorter shoots. Seedlings subjected to 65 W also had lower instantaneous rates of CO₂ assimilation, but higher instantaneous water-use efficiencies following irrigation, suggesting that koa responds to water deficits by decreasing water loss via reduced stomatal conductance. Seedlings subjected to 0P had similar rates of CO₂ assimilation relative to those grown with adequate P, suggesting that koa is able to employ strategies to avoid physiological impairment from conditions of inadequate P. Future research should assess whether subjecting koa seedlings to reduced water before planting on water-limited sites cues increased drought resistance and whether uptake and storage of P by seedlings in the nursery better supports growth following outplanting, particularly on sites with anticipated low plant-available water.

     

Survival and growth of drought hardened <Emphasis Type="Italic">Eucalyptus pilularis</Emphasis> Sm. seedlings and vegetative cuttings

Forestry requires low mortality of transplanted seedlings. Mortality shortly after planting is often associated with inadequate hydration of transplants. Seedlings can be hardened to the drought conditions they may experience after transplanting by exposing them to controlled drought conditions in the nursery. Eucalyptus pilularis Sm. seedlings were drought hardened by providing nil (severe treatment) or half (mild treatment) the daily irrigation routinely received (control treatment) for up to two non-consecutive days per week during the last 4 weeks of growth in the nursery. Drought hardening reduced stem diameter, seedling leaf area, leaf area per root biomass and seedling quality measured by the Dickson quality index, but increased root:shoot ratio. Hardened seedlings had lower stomatal conductance and leaf water potential on the days they received less irrigation that the control treatment. Hardened seedlings had greater stomatal conductance and were less water stressed than seedlings experiencing drought for the first time indicating hardened seedlings had adjusted physiologically to drought. Survival after transplanting in the controlled drought environment in a glasshouse was enhanced by the hardening treatments. Non hardened seedlings that had had their upper leaves manually removed immediately prior to transplanting to reduce leaf area (top-clipped) had similar survival to hardened seedlings. Stomatal conductance and leaf water potential after transplanting were higher in hardened and top-clipped seedlings than unhardened control seedlings or vegetative cuttings. Survival in the field trial was over 95% for all treatments, possibly as rain fell within 4 days of planting and follow-up rain occurred in the subsequent weeks. Neither the hardened or top-clipped seedlings planted in the field trial had reduced growth, increased propensity to form double leaders or worse stem form than control seedlings when measured at age 3 years.     

Phosphorus Enhanced Establishment,Growth, Nutrient Uptake,and Productivity of Dalbergia sissoo Seedlings Maintained at Varying Soil Water Stress Levels in an Indian Arid Zone

We assessed interactive effects of varying levels of applied phosphorus fertilizer and water stress on growth, productivity, and mineral accumulation in container-grown Dalbergia sissoo L. seedlings. Height, collar diameter, leaf size and area, root volume and total biomass were reduced, and dry matter allocation to root was increased with increasing levels of soil water stress. The reduction was >32% in growth, >50% in leaf, and >77% in biomass when seedlings were grown with <50% of soil field capacity. Phosphorus application at the level of 10 mg kg^?1 soil enhanced stems and leaf biomass and nutrient accumulation at all irrigation levels, and thus tolerance to drought. Phosphorus responses to growth and biomass production increased with irrigation levels. Thus, 20 mg P kg^?1 soil is beneficial at sufficient soil water availability and a lower dose (i.e., 10 mg P kg^?1) is recommended under high soil water stress conditions to benefit growth and productivity of D. sissoo.     

Production and establishment techniques for the restoration of Nothofagus alessandrii,an endangered keystone species in a Mediterranean forest

Ruil (Nothofagus alessandrii) is an endangered keystone species from the Mediterranean climate zone of Chile. Ruil’s fragile state of conservation urges development of restoration programs, but specific protocols for nursery production and field establishment that ensure plant survival are largely unknown. Therefore, we examined the effect on nitrogen (N) fertilization and container size during nursery production in combination with the use of mesh shelters after outplanting on survival and growth during the first growing season in the field. First year outplanting survival of nursery-grown container seedlings was enhanced when seedlings were given nitrogen (N) during nursery production and deployed with mesh tree shelters in the field. The volume of the container had no effect on outplanting survival and growth. Increasing N from zero to 200 mg N L^?1 provided sufficient N levels, resulting in increased seedling height, root-collar diameter, shoot biomass, and total seedling N and phosphorous concentrations. Additional N provided by the 400 and 600 mg N L^?1 treatments underwent luxury consumption by the seedlings with no further benefits in field performance. Improved growth in the nursery, along with the use of mesh tree shelters after outplanting, especially during the typical summer drought, may be responsible for increased survival during the first growing season. Increasing the performance of nursery-grown ruil seedlings is essential to restoring this endangered, vulnerable, and foundation species within the highly biodiverse, yet seriously threatened endemic Maulino Costero Forest of the Mediterranean climate of central Chile.

     

Effects of water stress on growth,dry matter allocation and water-use efficiency of a leguminous species, <Emphasis Type="Italic">Sophora davidii</Emphasis>

The adaptation responses to different water conditions and the drought tolerance of Sophora davidii seedlings were assessed in a greenhouse experiment. Two-month-old seedlings were subjected to the following water supplies for 95 days: 100, 80, 60, 40 and 20% of field water capacity. The seedlings at 100% FC had the greatest productivity, height, basal diameter, branch number, leaf number and leaf area. Water supply <80% FC was the threshold of drought-initiated negative effects on seedling growth, yield and physiological processes; these parameters were severely reduced at 20% FC, however, there was no plant death during the experiment. Moreover, water stress decreased leaf relative water content, specific leaf area, leaf area ratio, and water-use efficiency (WUE), whereas it increased the biomass allocation to roots, which resulted in a higher root:stem mass ratio under drought. The S. davidii seedlings tolerated drought by maintaining high leaf relative water content and by reducing branching and leaf expansion. However, low productivity and WUE at 20% FC suggested that seedlings did not produce high biomass under severe drought. Therefore, prior to introducing S. davidii in forestation efforts, a water supply >40% FC is recommended for seedlings to maintain growth and productivity. These results provide insights into limitations and opportunities for establishment of S. davidii in arid regions.     

Interacting effects of irradiance and water stress on dry weight and biomass partitioning in Fagus sylvatica seedlings

Abstract

Actual climate models for central Europe predict prolonged summer droughts. Knowledge on how the interaction between light and water availability affects regeneration will hence be of major importance. In an experiment carried out under controlled conditions, newly emerged beech seedlings were grown in pots with sand during 54 days. Three treatments applying three different light levels (2, 9 and 43% relative light intensity) were combined with two soil water treatments (control and drought). At the end of the experiment, seedlings were separated into leaves, stem and root and the seedlings’ dry mass, leaf area and stem length was determined. Low irradiance (2%) had a strong negative effect on dry weights of seedling components, leaf area and specific leaf mass. Drought clearly affected biomass partitioning for seedlings at high irradiance levels (43%). An interaction between irradiance and drought on biomass partitioning in beech seedlings was observed at medium irradiance level (9%). Within a wide range of light levels in the forests, both light and drought may affect biomass partitioning in young seedlings.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Organic amendment and inorganic fertilization affect soil properties and quality of Larix olgensis bareroot stock

Bareroot Changbai larch (Larix olgensis Henry.) seedlings were reared with inorganic fertilizer (nitrogen (N):phosphorus (P) = 1:1, W/W) applied at a rate of 100 (F100) or 200 kg N ha⁻¹ (F200) with (+) or without (−) chicken manure as a soil amendment (O) in north-eastern China. An unfertilized control treatment was included. Inorganic and organic fertilizer treatments tended to increase soil ammonium, nitrate, available P, total P, organic carbon content and electrical conductivity, and biomass and N concentration in seedlings. Organic amendment improved first order lateral root number, tap root length, fine root morphology (length, surface area, volume) in seedlings, while the F100 treatment increased N accumulation in needles and stems compared to the F200 treatment, on average. Most fertilizer treatments tended to increase P content in combined stems and roots, but F200 − O and F100 + O treatments diluted P in needles. Organic amendment combined with inorganic fertilizer at a rate of 100 kg N and P ha⁻¹ is recommended to improve seedling growth and N reserves in woody tissues.     

Effects of a deep container on morpho-functional characteristics and root colonization in Quercus suber L. seedlings for reforestation in Mediterranean climate

In the last decades, reforestation and afforestation programs are being carried out mainly with containerized seedlings. Container design determines the morphological and physiological characteristics of seedlings. However, container characteristics are often the same for plant species with very different growth strategies. The most commonly used nursery containers are relatively shallow and limit tap root growth; consequently, species relying on the early development of a long tap root to escape drought, such as those of the Quercus genus, might need to be cultivated in deep containers. The aim of this paper was to compare the morphological and physiological characteristics of Quercus suber L. seedlings cultivated in shallow containers (CCS-18, depth 18 cm) with seedlings cultivated in deep containers (CCL-30, depth 30 cm). Both container types used were made of high-density polyethylene, cylindrical in shape, open-bottomed, with a diameter of 5 cm, two kinds of vertical ribs on the inside wall showing a cultivation density of 318 seedlings/m². At the end of nursery culture, the seedlings cultivated in the CCL-30 deep container presented a longer tap root, higher shoot and root biomass and higher Dickson Quality Index (DQI). Moreover, the CCL-30 seedlings showed a higher root growth capacity (RGC), they reached deep substrate layers faster and they presented higher root hydraulic conductance. These morpho-functional advantages improved the CCL-30 seedling water status, which was expressed by higher stomatal conductance during an imposed drought period.     

Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water

Production and biomass allocation patterns, the growth rates of aboveground biomass, and crown traits were examined in saplings of the deciduous Quercus faginea and the evergreen Q. ilex to determine whether differences in these traits might account for the greater mortality during periods of drought undergone by Q. faginea. Strong differences were observed in almost all the traits analyzed, which suggests that the two species use different strategies to cope with the main limiting factors for woody seedling establishment in Mediterranean environments: excess light and low water availability. In Q. faginea, sapling design seems to be oriented to maximize light capture and, hence, leaf productivity during the short life span of the leaf biomass. Thus, the seedlings of Q. faginea showed crown traits that permit self-shading to be minimized: longer shoots with more spaced leaves that result in lower leaf area index than in Q. ilex. In addition, the larger area per unit leaf biomass in Q. faginea leads to a larger interceptive leaf area per unit plant mass and to higher light capture. These characteristics imply higher investments in woody tissues (SWR) that permit the plants to support a wide canopy and facilitate water transport to meet the strong transpiratory demands of a canopy with such characteristics. By contrast, in Q. ilex, saplings are apparently designed to guarantee leaf survival against temperature extremes and photoinhibition through avoidance of excessive radiation.     

Interaction of drought and elevated CO2 concentration on photosynthetic down-regulation and susceptibility to photoinhibition in Japanese white birch seedlings grown with limited N availability

The interaction of drought and elevated carbon dioxide concentration ([CO(2)]) on carboxylation capacity of Rubisco (V(cmax)) and susceptibility to photoinhibition may be an important determinant of plant responses to seasonal fluctuations in precipitation in an anticipated elevated [CO(2)] environment. Japanese white birch (Betula platyphylla var. japonica) leaves that developed wholly during a period of drought showed an increase in leaf nitrogen and a decrease in leaf carbohydrates that could ameliorate photosynthetic down-regulation, defined as a decrease in V(cmax) in response to elevated [CO(2)]. Photochemical quenching (q(P)) was decreased by elevated [CO(2)] but increased by drought when compared at a given intercellular [CO(2)] (C(i)), indicating that elevated [CO(2)] could increase the risk of photoinhibition, whereas long-term drought could alleviate the risk of photoinhibition. However, only a small variation in q(P) was measured among seedlings in the various water availability x [CO(2)] treatment combinations, consistent with the small treatment differences in chronic photoinhibition among the seedlings, as indicated by the ratio of variable to maximum chlorophyll fluorescence after overnight dark-adaptation. Our results suggest that the offsetting responses-reduced V(cmax) plus increased C(i) at elevated [CO(2)] and increased V(cmax) plus reduced C(i) under drought conditions-resulted in a narrow range of susceptibility to photoinhibition at the growth [CO(2)] in Japanese white birch seedlings grown in various water availability x [CO(2)] treatment combinations.     

Are marginal beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) provenances a source for drought tolerant ecotypes?

Due to the expected increases of number and intensity of summer droughts in Central Europe the identification of drought tolerant ecotypes becomes more important in future forestry. A common garden experiment with seedlings of Fagus sylvatica provenances from the center (Germany) and eastern margin (Poland) of the species’ distribution range was conducted. Responses of morphological, physiological, chemical and growth parameters to three drought treatments were analyzed. Relative growth rates of the marginal provenance were lower as compared to the central provenance. The marginal seedlings showed a tendency to higher total biomasses because of higher seed masses. In both provenances drought decreased biomass production and root/shoot ratio which was lower in the central provenance. A lower specific root area of the marginal provenance indicated a better adaptation to low xylem water potentials. Under moderate drought, lower leaf δ¹³C signatures may indicate lower stomatal limitation (or a reduced rate of CO₂ assimilation) in the marginal provenance. We conclude that marginal beech provenances may exhibit a better drought adaptation.     

落叶松水浸液对胡桃楸幼苗生长的影响

A study was conducted to detect the effect of water extracts from different parts such as root, bark, branch and leaf, of adult larch, Larix gmelini, trees on growth of Manchurian walnut, Juglans mandshudca, seedlings and the allelopathy between the two tree species. Four concentrations （100 g. kg i, 50 g. kg^-1, 25 g. kg^-1 and 12.5 g. kg^-1） were prepared for each kind of extracts. Result showed that the water extracts with low and moderate concentrations accelerated the growth of collar diameter and increased biomass and root/shoot ratio of walnut seedlings. The water extracts from branches and barks with low and moderate concentrations accelerated the height growth of the seedlings, while those from leaves and roots slightly decreased the height growth of seedlings. The fact that application of water extracts of larch improved the growth of Manchurian walnut attributes possibly to the allelopathy between the two tree species.     

Effects of arbuscular mycorrhizal fungi on the drought tolerance of Cyclobalanopsis glauca seedlings under greenhouse conditions

Cyclobalanopsis glauca is an important afforestation tree species that is widely used for revegetating the karst region of southwest China. Vegetation in this region is regularly commonly subjected to drought stress because of the geology and water shortages. Here, we investigated the influence of two arbuscular mycorrhizal fungi (AMF) Glomus mosseae and Glomus intraradices on the drought tolerance of C. glauca seedlings under greenhouse conditions. AMF-treated and non-AMF-treated C. glauca seedlings were maintained under two different water regimes (well watered: 80 % field capacity; drought stress: 40 % field capacity) for 90 days. The AMF colonization rate was higher under well-watered conditions compared to drought stress conditions. The growth and physiological performance of C. glauca seedlings were significantly affected by drought stress. Under drought stress conditions, mycorrhizal seedlings had greater height, base diameter, leaf area, and biomass compared to non-mycorrhizal seedlings. In addition, under drought conditions, AMF-inoculated seedlings had greater superoxide dismutase and peroxidase activity, higher soluble sugar content, and lower proline content compared to non-inoculated seedlings. Furthermore, AMF colonization increased the phosphorus and potassium content of seedling shoots under both well-watered and drought stress conditions. Therefore, AMF colonization enhanced the drought tolerance of C. glauca seedlings by improving growth performance, nutrient content, the quantity of osmotic adjustment compounds, and antioxidant enzyme activity. The results indicate that AMF are of potential use for the restoration of vegetation in the karst region of southwest China.     

Carbon dioxide enrichment improves growth, water relations and survival of droughted honey mesquite (Prosopis glandulosa) seedlings

Low water availability reduces the establishment of the invasive shrub Prosopis on some grasslands. Water deficit survival and traits that may contribute to the postponement or tolerance of plant dehydration were measured on seedlings of P. glandulosa Torr. var. glandulosa (honey mesquite) grown at CO(2) concentrations of 370 (ambient), 710, and 1050 micro mol mol(-1). Because elevated CO(2) decreases stomatal conductance, the number of seedlings per container in the elevated CO(2) treatments was increased to ensure that soil water content was depleted at similar rates in all treatments. Seedlings grown at elevated CO(2) had a greater root biomass and a higher ratio of lateral root to total root biomass than those grown at ambient CO(2) concentration; however, these seedlings also shed more leaves and retained smaller leaves. These changes, together with a reduced transpiration/leaf area ratio at elevated CO(2), may have contributed to a slight increase in xylem pressure potentials of seedlings in the 1050 micro mol mol(-1) CO(2) treatment during the first 37 days of growth (0.26 to 0.40 MPa). Osmotic potential was not affected by CO(2) treatment. Increasing the CO(2) concentration to 710 and 1050 micro mol mol(-1) more than doubled the percentage survival of seedlings from which water was withheld for 65 days. Carbon dioxide enrichment significantly increased survival from 0% to about 40% among seedlings that experienced the lowest soil water content. By increasing seedling survival of drought, rising atmospheric CO(2) concentration may increase abundance of P. glandulosa on grasslands where low water availability limits its establishment.     

Efficiency of planting hybrid larch F1 ( Larix gmelinii var. japonica × L. kaempferi ) at different planting densities and using seedlings or rooted cuttings

The efficiency of planting a hybrid larch F₁ at two planting densities and of using seedlings of hybrid larch F₁ or rooted cuttings of Gream, the registered variety, was investigated. Significantly more time was required to move from one planting spot to the next in plots with 625 seedlings/ha than in plots with 1,333 seedlings/ha, but there was no significant difference between seedlings and rooted cuttings. Approximately 18% more time was needed to plant a single planting stock at the low planting density than at the high density. The need to prune roots or enlarge a planting hole occurred more often for rooted cuttings than for seedlings, and 17% more time was needed to plant rooted cuttings of Gream than seedlings.     

不同土壤水分胁迫格局对印度沙漠中的黄檀种苗营养吸收和生物量的影响

1998年7月,利用非称重式蒸渗池种植单一种源的一年生黄檀种苗,研究在印度沙漠地区培养黄檀种苗的合理灌水技术参数。当各处理（W1、W2、W3、W4）的土壤水分含量分别降低到7.56%、5.79%、4.44%和3.23%时,通过灌溉使苗木生长保持在一定的土壤的水分状况,如36.2mm（W1）、26.5mm（W2）、20.2mm（W3）和18.1mm（W4）。结果表明,在36.2mm（W1）水平时,种苗的株高、冠径、叶数和叶面积达到最大值（p〈0.01）。在W1和W2处理中,虽然上述参数没有明显差异,但在W2处理中,种苗的每升水分利用率的生物量最大。在W3、W4和W5灌溉水平下,不利于提高种苗的株高、生物量和营养积累。在W2水平以下（5.79%）,土壤水分有效率能提高根系生物量占总生物量的百分比。但在W3和W4处理中,叶干生物量百分比下降,同时在W5处理中,茎干生物量百分比下降。在W5处理中,土壤水势达到-196Mpa,种苗才可以成活。在W3和W4处理中,土壤水分有效率的限额影响黄檀种苗的生长和生物量。在W2处理中,土壤水分有效利用率最高,种苗的生长和生物量达到最高值。因此,在壤砂土条件下,通过灌溉维持幼苗土壤水分含量在5.79%以上时,可获得较好的黄檀种苗的生长和生物量产量。     

Seasonal evolution of water status after outplanting of two provenances of Holm oak nursery seedlings

Forest restoration programs using Holm oak (Quercus ilex ssp. ballota [Desf.] Samp.) have had limited success. The effect of plant provenance on plantation success is uncertain, although some previous studies suggest that some provenances may be better able to tolerate stress. We studied the tolerance to drought in seedlings from two Spanish provenances of Holm oak before and after outplanting. One provenance was from a continental climate with cold winters (GR) and the other was from a xeric climate (HU). Seedlings were subjected to a water stress test in the nursery during the summer and survival was visually assessed after 2?weeks. In addition, 35 healthy seedlings of each provenance that were not subjected to the water stress tests were used for outplanting experiment. In these plants the seasonal changes in water potential at dawn (Ψ), specific leaf area (SLA), cuticular transpiration (E_c), and loss of xylem hydraulic conductance of twigs (PLC) were measured over 18?months. After the water stress test in summer, mortality was 44.3?% for GR seedlings and 12.6?% for HU seedlings. In addition there were differences between the two provenances in plant water status after planting. The HU provenance had a better water status and was more water conservative in the summer (higher Ψ, lower E_c, lower PLC), but not in the winter. The different drought tolerance and water relations parameters of these two provenances indicate that provenance should be considered in forest restoration and conservation programs involving Holm oak.     

Growth, physiology, and nutrient retranslocation in nitrogen-15 fertilized Quercus rubra seedlings

We evaluated gas exchange, chlorophyll index (CI) using SPAD-502 chlorophyll meter, and leaf nutritional responses in one-year-old northern red oak (Quercus rubra L.) container seedlings transplanted into control (unfertilized) or fertilized (0.86 g N plant^?1) sand culture and grown in a greenhouse for 90 days. We labeled current nitrogen (N) uptake with (¹⁵NH₄)₂SO₄ and directly quantified proportional contributions of N derived from fertilizer (NF) compared with retranslocation or N derived from plant (NP) in leaf growth of red oak seedlings. NF met a greater N demand in leaf growth of fertilized plants while unfertilized seedlings relied entirely on NP for their leaf growth. Fertilization increased leaf dry mass by 67% and new stem dry mass by 253% 90 days after transplanting compared to control seedlings. Specific leaf area (SLA) was not significantly altered but CI increased 90 days after transplanting. Higher leaf N concentration and content in fertilized compared with control seedlings was linked to greater chlorophyll concentrations in the former plants. The higher coefficient of determination (r ² = 0.72) noted between leaf N concentrations and CI suggests that the SPAD meter could be a useful tool for assessing leaf N status in northern red oak seedlings. Fertilized seedlings exhibited higher net assimilation, stomatal conductance, and transpiration rates compared with controls. Increased seedling growth in response to fertilization was related to maintenance of higher gas exchange and greater nutrient uptake, which could improve outplanting success.     

Morphological response of Vitex negundo var. heterophylla and Ziziphus jujuba var. spinosa to the combined impact of drought and shade

Light and soil moisture availability are two important abiotic factors influencing plant growth in an agroforestry system. Different soil moisture and light treatments were applied to examine the combined impact of drought and shade on the morphological plasticity of Vitex negundo var. heterophylla (Chinese chastetree) and Ziziphus jujuba var. spinosa (Spine jujube). We found that the interaction of light and soil moisture was orthogonal in the two species. V. negundo captured irradiance efficiently with relatively long petiole and petiolule, while Z. jujuba maintained higher branches to absorb light than V. negundo. Compared to the seedlings under full sunlight, the palmately compound leaves of V. negundo under low light showed larger specific leaf area (SLA), lower ratio of leaflet length to width and higher leaf mass ratio (leaf biomass to total biomass); in contrast, the simple leaves of Z. jujuba under low light showed larger SLA and ratio of leaf length to petiole length. In both species, drought reduced the branch number, mean internode length of stem, and increased root mass ratio (root biomass to total biomass), but leaf morphology showed little variation to the decreased soil moisture. Between the two species, V. negundo is more shade-tolerant, while Z. jujuba is more drought-tolerant. In an alley cropping system, the spacing can be smaller in the V. negundo-crop system than the Z. jujuba-crop system when soil moisture is adequate. However, the V. negundo-crop system should be managed with more caution under the soil moisture limited conditions.