Approaches to modelling the uptake of water and nutrients in agroforestry systems

This review presents information about root systems of crops and trees and describes approaches that have been used to model uptake of water and nutrients in crops that may have application to agroforestry systems. Only a few measurements of the distribution of tree roots in agroforestry systems have been published and these are predominantly in alley cropping systems with young trees. Therefore, a major limitation to developing water and nutrient uptake models for trees is the lack of adequate measurements and conceptual models for describing the distribution of roots spatially and temporally. Most process-based modelling approaches to water and nutrient uptake integrate the activities of a single root over the whole root system. Several difficulties can be foreseen with applying these approaches to roots of older trees including the presence of mycorrhizal associations so that the root surface is not the site of uptake, the uncertainty as to whether all tree roots are active in taking up water and nutrients, and the fact that, unlike annual crops, trees have substantial reserves of nutrients that can be mobilised to support growth so that the notion of a plant demand regulating uptake may prove difficult to define. The review concludes that a programme of experimental measurements is required together with modelling using approaches both in which roots are implicit, and in which process-based models with roots allow competitive ability to be assessed.     

Soil aeration and growth of forest trees (review article)

Many forest trees exhibit reduced growth or are killed when the soil is low in oxygen. Anaerobic soil conditions are associated with flooding or compaction of soil but also occur commonly in soils of heavy texture. Reductions in height growth, leaf growth, cambial growth and reproductive growth of trees growing on poorly aerated soils are well documented. The amount of growth reduction varies widely among species and duration of anaerobic soil conditions during the growing season. Inhibition of growth is preceded by changes in physiological processes, including food, water, hormone, and mineral relations. Some species can adapt to soil anaerobiosis by (1) producing hypertrophied lenticels which assist in aeration of the stem and release of toxic compounds, and (2) growing new roots to replace loss of original roots under anaerobic conditions. The replacement roots assist in absorption of water and mineral nutrients and in oxidizing the rhizosphere and detoxifying soil toxins. Ethylene, together with other compounds, appears to play an important causal role in morphological adaptations to soil anaerobiosis. Some forest trees can also adapt metabolically to poor soil aeration.     

A model analysis of the influence of root and foliage allocation on forest production and competition between trees

A general model was constructed relating forest growth to nitrogen uptake and the partitioning of biomass among leaves, fine roots and woody tissues. The model was used to assess the influence of the allocation pattern on stand wood production, individual tree growth and nutrient cycling for even-aged conifer stands with adequate water, but suboptimal nitrogen. Stand wood production was maximized by quite low allocation to roots for specified amounts of plant-available nitrogen. However, the wood production of the individual was maximized by higher allocation to roots, because large root systems enhanced the ability of individuals to compete for nutrients. The optimal fine root allocation for a competing individual was less than 5% of total production for adequate nitrogen, but rose to 30% as nitrogen became more limiting, in general agreement with observed allocation patterns for fertilized versus non-fertilized forests. The high allocation to roots predicted for competitors may also enhance long-term productivity by decreasing nutrient losses from the ecosystem. Although collective, short-term stand wood production could be increased by shifting growth from roots to stems, this strategy may increase nutrient losses, ultimately decreasing productivity.     

Effects of compaction and simulated root channels in the subsoil on root development, water uptake and growth of radiata pine

Effects of subsoil compaction and simulated root channels (perforations) through the compacted layer on root growth, water uptake, foliar nutrient concentration and growth of radiata pine (Pinus radiata D. Don) were studied in a field experiment where a range of treatments were applied in reconstituted soil profiles. Subsoil compaction adversely affected root penetration in deeper parts of the soil and consequently caused greater water stress in trees. However, the effect of compaction was largely overcome when the subsoil was perforated to render 0.2% of the soil volume into vertical channels. Roots showed a remarkable ability to reach the points of low penetration strength and to travel through them to deeper parts of the profile. Perforations through compacted soil layers at a relatively low frequency may be a practical solution to allow root development into deeper parts of the soil and allow greater soil water exploration by roots.     

黄土高原刺槐根系垂直分布特征研究

在中国西北黄土高原地区，水分是树木生长发育的主要限制因子。根系分布特征由于反映了树木对环境条件的利用程度而具有更加特殊的生态意义。本研究选择阳坡和阴坡不同立地上年龄一致的刺槐林调查了根系分布特征。根系垂直分布特征的调查结果表明，在所有立地上，根系生物量随着深度的增加而降低，其中细根的分布深度大于粗根的分布深度。方差分析结果表明：不同立地上不同径级根系的分布特征也有明显的差别，粗根是差异存在的主要原因，阴坡立地上的根系生物量，特别是细根生物量大于阳坡立地上的。对根系消弱系数的分析结果表明，阴坡立地上的根系消弱系数大于0．982，而阳坡立地上的根系消弱系数小于0．982，说明阴坡市地上刺槐根系的生物量在深层土壤中的分布相对量更大一些。其中细根的根系消弱系数大于粗根的，这种根系分布特征有利于根系对深层土壤水分养分的吸收利用，进而促进树木地上部分的生长发育。图3表3参15。     

Responses of carbon gain and growth of Pinus radiata stands to thinning and fertilizing

Thinning of forest stands is widely carried out to minimize the slowing of growth of individual stems that follows from increasing competition among trees as they become bigger. After thinning, there is an increase in the growth rate of remaining trees because of an increase in the availability of resources per tree. Often, there is also an increase in foliar efficiency (biomass increase/foliage amount). On sites where mineral nutrient supply is limiting, fertilizers may be applied, often in association with thinning, to boost productivity. Growth responses to fertilizer application depend on an adequate supply of other resources, but also involve nonlinear interactions among mineral nutrients and between nutrients and other growth-limiting environmental factors. The effects of thinning and fertilizing on the carbon gain and growth responses of Pinus radiata D. Don to availability of resources (light, mineral nutrients and water) and to changes in the canopy are discussed.     

Tree-grass relationships in open eucalypt woodlands of northeastern Australia: influence of trees on pasture productivity, forage quality and species distribution

Pasture yield, quality and species distributions were compared between zones around live and killed eucalypt trees at two woodland sites in northeast Queensland which differed markedly in soil fertility. Trees affected pasture quality and yield on an individual tree basis: N concentration and dry matter digestibility tended to be higher under trees than in inter-tree areas at both sites and pasture yields declined with distance from killed trees at the lower fertility site. However, the distribution of individual species did not vary markedly with distance from trees. Trees also affected pasture yield on a woodland basis: yields were greater where the trees were killed than under intact woodland. Soil under trees had higher levels of organic carbon and greater litter cover than soil in inter-tree areas. However, pasture yields did not generally reflect this fertility gradient since growth was limited by moisture availability due to drought conditions during the study period. Pasture N concentration was higher under trees that in inter-tree areas since plants under trees produced a similar amount of biomass as plants in inter-tree areas, but had access to higher nutrient levels. Trees appeared to have a greater effect on soil nutrient availability at the low fertility site. Live trees depressed pasture yields to a lesser degree at the low fertility site, demonstrating that the effects of trees on soil water availability (on a woodland scale) are less important when soil nutrients are more limiting to growth. These results indicate that, while removing trees may enhance pasture productivity, this benefit may be offset by a reduction in pasture quality. Given the beneficial effect of trees on soil nutrients, tree removal may also have longer term implications for soil nutrient dynamics. This revised version was published online in August 2006 with corrections to the Cover Date.     

Responses of Picea, Pinus and Pseudotsuga roots to heterogeneous nutrient distribution in soil

The spatial distribution of plant-available mineral nutrients in forest soils is often highly heterogeneous. To test the hypothesis that local nutrient enrichment of soil leads to increased root proliferation in the nutrient-rich soil zone, we studied the effects of nutrient enrichment on the growth and nutrient concentrations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) roots. Three-year-old seedlings were grown for 9 months in split-root containers filled with nutrient-poor forest mineral soil, with one side supplemented with additional mineral nutrients. Root dry weight and root length in Scots pine and Norway spruce were increased in the nutrient-supplemented soil compared with the nonsupplemented side, whereas root growth in Douglas-fir was unaffected by nutrient enrichment. Of the three species examined, Norway spruce exhibited the highest root and shoot growth and the highest nutrient demand. Specific root length (m g(-1)) and the number of root tips per unit root length were not affected by local nutrient addition in any of the species. Despite increased root growth in Norway spruce and Scots pine in nutrient-supplemented soil, their root systems contained similar nutrient concentrations on both sides of the split-root container. Thus, coniferous trees may respond to local nutrient supply by increased root proliferation, but the response varies depending on the species, and may only occur when trees are nutrient deficient. As a response to local nutrient enrichment, increases in root dry matter or root length may be better indicators of pre-existing nutrient deficiencies in conifers than increases in root nutrient concentrations.     

Belowground interactions for water between trees and grasses in a temperate semiarid agroforestry system

A fundamental hypothesis of agroforestry is the complementary use of soil resources. However, productivity of many agroforestry systems has been lower than expected due to net competition for water, highlighting the need for a mechanistic understanding of belowground interactions. The goal of this study was to examine root–root interactions for water in a temperate semiarid agroforestry system, based on ponderosa pines and a Patagonian grass. The hypotheses were: (a) A greater proportion of water uptake by pines is from deeper soil layers when they are growing with grasses than when they are growing alone; (b) Growth of grasses is improved by the use of water hydraulically lifted by pines. We used stable isotopes of O to analyze water sources of plants, and we measured sapflow direction in pine roots and continuous soil water content with a very sensitive system. We also installed barriers to isolate the roots of a set of grasses from pine roots, in which we measured water status, relative growth and water sources, comparing to control plants. The results indicated that pines and grasses show some complementary in the use of soil water, and that pines in agroforestry systems use less shallow water than pines in monoculture. We found evidence of hydraulic lift, but contradicting results were obtained comparing growth and isotope results of the root isolation experiment. Therefore, we could not reject nor accept that grasses use water that is hydraulically lifted by the pines, or that this results in a positive effect on grass growth. This information may contribute to understand the complex and variable belowground interactions in temperate agroforestry.     

Coarse root elongation rate estimates for interior Douglas-fir

Accurate estimates of root growth rates are important for root system modeling, and the spread of root systems may be an important determinant of belowground site occupancy. Estimating root system growth rates is complicated because missing, discontinuous, and false annual growth rings make root cross sections difficult to age. These irregularities can occur even in roots of dominant conifers with rare or absent stem growth ring abnormalities. Incomplete rings were noted in the root growth rings of nine co-dominant interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees. Coarse root (> 1.0-cm diameter) elongation rates were estimated by fitting a geometric mean regression line to ring count and lateral distance data. In all nine roots examined, the geometric mean regression slope was well within the range of the 95% confidence interval for the ordinary least squares regression of lateral distance versus age, suggesting that measurement error may have been negligible. Coarse root elongation rates (which ranged from 2.8 to 15.3 cm year(-1) and averaged 7.4 cm year(-1)) in the interior Douglas-fir trees studied were much lower than those reported by others. This discrepancy may be a result of limited soil water availability, soil heterogeneity (both soil water content and soil texture were highly variable across short distances) and fragmentation of belowground growing space.     

Eucalyptus urophylla root growth, stem sprouting and nutrient supply from the roots and soil

Lack of information concerning root growth of trees limits our knowledge of plant development and fertilizer response. The objective of this work was to study root growth dynamics of an E. urophylla forest after harvesting and the supply of nutrients from the roots and the soil to the new sprouts originating from the stumps. About 7-year-old eucalypt trees were felled and the sprouts and roots were sampled at 0, 60, 120, 180, 240, 330 days after harvesting. The roots were separated into fine roots (<1 mm), medium roots (1–3 mm), coarse roots (>3 mm), and taproot. Nutrient supply to sprouts from the old roots and the soil was calculated based on the change in nutrient content of the roots with time and accumulation of nutrients in the sprouts. Fine, medium and coarse root biomass increased with time after harvesting. However, the increase was more pronounced with fine roots. Between harvesting and day 60 of the new growth, all nutrients allocated to the sprouts, excluding potassium, were supplied by the soil. K was the nutrient most dependent on root reserves for the initial growth of sprouts. The contribution of the old roots to N, P, Ca, and Mg accumulation in the sprouts increased between day 60 and 120. At 330 days after harvesting, about 9.2, 23.9, and 12.6% of the N, K, and Mg, respectively, that had accumulated in the sprouts were supplied by the roots, while all P and Ca were supplied by the soil.     

Single-tree influences on soil properties in agroforestry: lessons from natural forest and savanna ecosystems

Climate, organisms, topographic relief, and parent material interacting through time are the dominant factors that control processes of soil formation and determine soil properties. In both forest and savanna ecosystems, trees affect soil properties through several pathways. Trees alter inputs to the soil system by increasing capture of wetfall and dryfall and by adding to soil N via N₂-fixation. They affect the morphology and chemical conditions of the soil as a result of the characteristics of above- and below-ground litter inputs. The chemical and physical nature of leaf, bark, branch, and roots alter decomposition and nutrient availability via controls on soil water and the soil fauna involved in litter breakdown. Extensive lateral root systems scavenge soil nutrients and redistribute them beneath tree canopies. In general, trees represent both conduits through which nutrients cycle and sites for the accumulation of nutrients within a landscape. From an ecological perspective, the soil patches found beneath tree canopies are important local and regional nutrient reserves that influence community structure and ecosystem function. Understanding species-specific differences in tree-soil interactions has important and immediate interest to farmers and agroforesters concerned with maintaining or increasing site productivity. Lessons from natural plant-soil systems provide a guide for predicting the direction and magnitude of tree influences on soil in agroforestry settings. The challenge for agroforesters is to determine under what conditions positive tree effects will accumulate simultaneously within active farming systems and which require rotation of cropping and forest fallows.     

Soil temperature and plant growth stage influence nitrogen uptake and amino acid concentration of apple during early spring growth

In spring, nitrogen (N) uptake by apple roots begins about 3 weeks after bud break. We used 1-year-old 'Fuji' Malus domestica Borkh on M26 bare-root apple trees to determine whether the onset of N uptake in spring is dependent solely on the growth stage of the plant or is a function of soil temperature. Five times during early season growth, N uptake and total amino acid concentration were measured in trees growing at aboveground day/night temperatures of 23/15 degrees C and belowground temperatures of 8, 12, 16 or 20 degrees C. We used (15NH4)(15NO3) to measure total N uptake and rate of uptake and found that both were significantly influenced by both soil temperature and plant growth stage. Rate of uptake of 15N increased with increasing soil temperature and changed with plant growth stage. Before bud break, 15N was not detected in trees growing in the 8 degrees C soil treatment, whereas 15N uptake increased with increasing soil temperatures between 12 and 20 degrees C. Ten days after bud break, 15N was still not detected in trees growing in the 8 degrees C soil treatment, although total 15N uptake and uptake rate continued to increase with increasing soil temperatures between 12 and 20 degrees C. Twenty-one days after bud break, trees in all temperature treatments were able to acquire 15N from the soil, although the amount of uptake increased with increasing soil temperature. Distribution of 15N in trees changed as plants grew. Most of the 15N absorbed by trees before bud break (approximately 5% of 15N supplied per tree) remained in the roots. Forty-six days after bud break, approximately one-third of the 15N absorbed by the trees in the 12-20 degrees C soil temperature treatments remained in the roots, whereas the shank, stem and new growth contained about two-thirds of the 15N taken up by the roots. Total amino acid concentration and distribution of amino acids in trees changed with plant growth stage, but only the amino acid concentration in new growth and roots was affected by soil temperature. We conclude that a combination of low soil temperature and plant developmental stage influences the ability of apple trees to take up and use N from the soil in the spring. Thus, early fertilizer application in the spring when soil temperatures are low or when the aboveground portion of the tree is not actively growing may be ineffective in promoting N uptake.     

Vertical and horizontal water redistribution in Norway spruce (Picea abies) roots in the Moravian Upland

Hydraulic redistribution (HR) by roots of large Norway spruce (Picea abies (L.) Karst.) trees was investigated by means of sap flow measurements made with the heat field deformation method. Irrigation was applied to a limited portion of the root system to steepen gradients of water potential in the soil and thus enhance rates of HR. On completion of the sap flow measurements, and to aid in their interpretation, the structure of the root system of seven of the investigated trees was exposed to a depth of 30 cm with a supersonic air-stream (air-spade). Before irrigation, vertical redistribution of water was observed in large coarse roots and some adjacent small lateral roots. Immediately after localized irrigation, horizontal redistribution of water from watered roots to dry roots via the stem base was demonstrated. The amount of horizontal distribution depended on the position of the receiving roots relative to the watered roots and the absorbing area of the watered root. No redistribution from watered roots via dry soil to roots of neighboring trees was detected. Responses of sap flow to localized irrigation were more pronounced in small lateral roots than in large branching roots where release and uptake of water are integrated. Sap flow measurements with multi-point sensors along radii in large lateral roots demonstrated water extraction from different soil horizons. We conclude that synchronous measurements of sap flow in both small and large lateral roots are needed to study water absorption and transport in tree root systems.     

Water stress, photosynthesis and early growth patterns of cuttings of three Populus clones

Photosynthetic attributes, leaf area and early root growth patterns were studied in three Populus clones to identify traits associated with superior growth potential on sites where water could be a limiting factor. It was found that early root growth and superior leaf area production were more closely related to growth potential than were photosynthetic capacity or carboxylation efficiency. A hybrid clone of Populus nigra var. charkowiensis (syn. P. nigra var. plantierensis) x P. nigra cv. 'Incrassata' (NE308) had more leaf area production and greater root system development in both wet and dry soil than did a P. trichocarpa clone (T6) and a P. balsamifera clone (B3). Despite greater above- and below-ground productivity, plants of clone NE308 had significantly lower photosynthetic capacity and carboxylation efficiency and a slightly higher CO(2) compensation point than plants of clones T6 and B3. Rapid early leaf and root growth appear to be key attributes associated with productivity in these clones regardless of soil water availability.     

Driving competitive and facilitative interactions in oak dehesas through management practices

Dehesas are extant multi-purpose agroforestry systems that consist of a mosaic of widely-spaced scattered oaks (Quercus ilex L.) combined with crops, pasture or shrubs. We aimed to clarify the role of trees in dehesas of CW Spain focussed on the analysis of tree-understorey interactions. Spatial variability of resources (light, soil moisture and fertility), microclimate, fine roots of both herbaceous plants and trees and forage yield was measured. Additionally, we compared the nutritional and physiological status, growth and acorn production of oaks in cropped (fodder crop), grazed (native grasses) and encroached (woody understorey) dehesa plots. Significant light interception by trees was limited to the close vicinity of the trees, with very low reduction away from them. Both microclimate and soil fertility improved significantly in the trees vicinity, irrespective of soil management. Soil moisture varied very few with distance from the trees, as a result of the extended root system of oaks. Root systems of trees and herbs did not overlap to a great extent. Crop production was higher beneath trees than beyond the trees in unfertilised plots and foliar nutrient content of oaks did not increase significantly with crop fertilisation, indicating that trees and crops hardly compete for nutrients. Moreover, trees benefited from the crop or pasture management: trees featured a significantly improved nutritional and physiological status, a faster growth and a higher fruit productivity than trees growing in encroached or forest plots.     

Soil-plant hydrology of indigenous and exotic trees in an Ethiopian montane forest

Fast-growing exotic trees are widely planted in the tropics to counteract deforestation; however, their patterns of water use could be detrimental to overall ecosystem productivity through their impact on ecosystem water budget. In a comparative field study on seasonal soil-plant water dynamics of two exotic species (Cupressus lusitanica Mill. and Eucalyptus globulus Labill.) and the indigenous Podocarpus falcatus (Thunb.) Mirb. in south Ethiopia, we combined a 2.5-year record for climate and soil water availability, natural-abundance oxygen isotope ratios (delta(18)O) of soil and xylem water, destructive root sampling and transpiration measurements. Soil was generally driest under C. lusitanica with its dense canopy and shallow root system, particularly following a relatively low-rainfall wet season, with the wettest soil under E. globulus. Wet season transpiration of C. lusitanica was twice that of the other species. In the dry season, P. falcatus and C. lusitanica reduced transpiration by a factor of six and two, respectively, whereas E. globulus showed a fivefold increase. In all species, there was a shift in water uptake to deeper soil layers as the dry season progressed, accompanied by relocation of live fine root biomass (LFR) of C. lusitanica and P. falcatus to deeper layers. Under P. falcatus, variability in soil matric potential, narrow delta(18)O depth gradients and high LFR indicated fast water redistribution. Subsoil water uptake was important only for E. globulus, which had low topsoil LFR and tap roots exploiting deep water. Although P. falcatus appeared better adapted to varying soil water availability than the exotic species, both conifers decreased growth substantially during dry weather. Growth of E. globulus was largely independent of topsoil water content, giving it the potential to cause substantial dry-season groundwater depletion.     

Family variation in nutritional and growth traits in Douglas-fir seedlings

Nitrogen (N) uptake and utilization in seedlings of six full-sib families of coastal Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) known to differ in growth rate were assessed at the whole plant and root levels. Seedlings were grown in soil or aeroponically with high and low nutrient availability. Consistent family differences in growth rate and N utilization index were observed in both soil and aeroponic culture, and high-ranking families by these measures also had greater net N uptake in soil culture. Two of the three families found to be fast-growing in long-term field trials exhibited faster growth, higher N utilization indices and greater net N uptake at the seedling stage. Mean family net influx of ammonium (NH4+) and efflux of nitrate (NO3-) in the high- and low-nutrient treatments were significantly correlated with measures of mean family biomass. The high-nutrient availability treatment increased mean net fluxes of NH4+ and NO3- in roots. These results indicate that efficiency of nutrient uptake and utilization contribute to higher growth rates of trees. Nutrient-related traits should be considered in tree breeding programs, as the indications are that assessments may be made at an early stage.     

环境因子对树木细根生物量、生产与周转的影响

细根在森林生态系统C平衡和养分循环中的重要作用已为大量研究所证实，树木有赖于细根吸收水分和养分，而细根对环境胁迫比较敏感，因此细根动态可指示环境变化，还可反映树木的健康状态，影响树木细根生产和周转的因子很多，本文在收集大量研究文献基础上，讨论了文献基础上，讨论了土壤养分，水分、pH值，温度等环境因子以及大气CO2增长对树木细根分布，生物量，生产和周转的影响，以期为我国开展细根生态学研究提供参考。