Andrew R. Gillespie

Interspecific interactions in temperate agroforestry

The ecological principles that define the competitive and complementary interactions among trees, crops, and fauna in agroforestry systems have received considerable research attention during the recent past. These principles have not yet, however, been adequately integrated and synthesized into an operational approach. This paper reviews the ecological and ecophysiological bases for interspecific interactions based on data from site-specific research and demonstration trials from temperate agroforestry systems, primarily from temperate North America. The review shows that information on ecological interactions in several temperate agroforestry systems is inadequate. It is recommended that the future research should focus on exploring new species and systems that have received little attention in the past. Priority research areas should include cultural practices and system designs to minimize interspecific competition and maximize environmental benefits such as improved water quality. Potential for genetic modification of components to increase productivity and reduce competition also needs to be explored. Process-oriented models may be used increasingly to predict resource-allocation patterns and possible benefits for a suite of site and species combinations.

Defining competition vectors in a temperate alley cropping system in the midwestern USA; 2. Competition for water.

A study was carried out in an alley cropping system in the Eastern Maize Belt of the United States with the objective of quantifying the competition for water between trees (black walnut (Juglans nigra L.) and red oak (Quercus rubra L.) and maize (Zea mays L.). Spatial and temporal variation in soil moisture, tree and maize rooting patterns, tree and maize water uptake, and maize leaf area expansion were determined in three treatments; ‘barrier’ (polyethylene root barriers separating maize and tree roots), ‘trench’ (trenching without a polyethylene barrier), and ‘no barrier’ (control). Significant temporal variation in soil moisture was observed in both black walnut and red oak alley cropping systems. The ‘barrier’ and ‘trench’ treatments resulted in higher soil moisture in the ‘maize alley’ and lower soil moisture in the ‘tree row’ than the ‘no barrier’ treatment. Uptake of water by maize was higher in the ‘barrier’ than the ‘no barrier’ treatment. However, tree water uptake was higher in the ‘no barrier’ treatment than in the ‘barrier’ treatment because tree roots utilized water from the ‘maize alley’. Maize leaf area was negatively impacted by water stress in the ‘no barrier’ treatment. Quantification of rooting patterns revealed that maize and tree roots were concentrated in the top 30 cm soil layer where water fluctuations were greatest. The ‘barrier’ and ‘trench’ treatments successfully eliminated the belowground root competition between trees and maize and resulted in greater leaf area and higher grain yields than the ‘no barrier’ treatment. Thus, competition for water rather than competition for light seems to be critical in defining the productivity and sustainability of this alley cropping system.

Defining competition vectors in a temperate alley cropping system in the midwestern USA; 3. Competition for nitrogen and litter decomposition dynamics.

An experiment was conducted in an 11-year-old black walnut (Juglans nigra L.), red oak (Quercus rubra L.), maize (Zea mays L.) alley cropping system in the midwestern USA to examine the extent of tree-crop competition for nitrogen and decomposition dynamics of tree leaves and fine roots. A below-ground polyethylene root barrier (1.2 m deep) isolated black walnut roots from maize alleys in half the number of plots providing two treatments viz. ‘barrier’ and ‘no barrier’. The percentage of N derived from fertilizer (%NDF) and fertilizer use efficiency (%UFN) were determined using 15N enriched fertilizer. Further, maize grain and stover biomass, tree leaf biomass, tissue N concentration, and N content were quantified in both treatments. The ‘barrier’ treatment resulted in a significantly greater grain (67.3% more) and stover (37.2% more) biomass than the ‘no barrier’ treatment. The %NDF in both grain and stover was higher in the ‘no barrier’ treatment as a result of competition from tree roots for water and mineralized N in soil. Maize plants growing in the ‘no barrier’ treatment had a lower %UFN than those in the ‘barrier’ treatment due to their smaller size and inability to take up fertilizer. Analysis of tree leaf and fine root decomposition patterns revealed faster release of N (39% over 15 days for black walnut and 17.7% for red oak) and P (30% over 15 days for both species) from roots compared to the leaves of both species. Following an early release of P (11.3% over 45 days), red oak leaves exhibited significant immobilization for the rest of the incubation period. The data indicate that competition for N from fertilizer is minimal since nutrient acquisition is not simultaneous among black walnut and maize. However, competition for mineralized N in soil can exist between black walnut and maize depending on water availability and competition. Tree leaves and fine roots can enhance soil nutrient pools through the addition of soil carbon and nutrients. Tree fine roots seem to play a more significant role in nutrient cycling within the alley cropping system because of their faster release of both N and P as compared to leaves. Selection of tree species and their phenology will impact the magnitude and rate of nutrient cycling.

Allelopathy in black walnut (Juglans nigra L.) alley cropping. II. Effects of juglone on hydroponically grown corn (Zea mays L.) and soybean (Glycine max L. Merr.) growth and physiology.

We conducted an experiment to investigate the effects of juglone (5-hydroxy-1, 4-napthoquinone) on the growth and physiology of hydroponically grown corn (Zea mays L.) and soybean (Glycine max L. Merr.) seedlings. Three different concentrations of juglone (10-6 M, 10-5 M, and 10-4 M) along with a control were applied. Within 3 days, juglone exhibited significant inhibitory effects on all measured variables including shoot and root relative growth rates (RGRs and RGRr), leaf photosynthesis (Pnet), transpiration (E), stomatal conductance (gs), and leaf and root respiration. In general, soybean was found to be more sensitive to juglone than corn. RGRr was the most inhibited variable for both species, and reductions of 86.5 and 99% were observed in corn and soybean, respectively, with 10-4 M juglone concentrations. Among the physiological variables measured, Pnet showed the greatest impact of toxicity though the other physiological parameters were also impacted. We conclude that both corn and soybean are sensitive to juglone and observed growth reductions in corn and soybean in black walnut alley cropping may partly be due to juglone phytotoxicity. Determination of actual phytotoxicity will require quantification of soil solution juglone levels, particularly in areas where soil solid-phase levels are high in close proximity to trees.

Defining competition vectors in a temperate alley cropping system in the midwestern USA: 1. Production physiology

With renewed interest in the use of ecologically-designed, sustainable agricultural systems for temperate regions of the world, agroforestry is being proposed as an alternative to intensive production of crops in monocultures. However, the knowledge-base for understanding and managing complex, multi-strata systems worldwide is limited, particularly so for temperate regions. We examined an alley cropping system in the midwestern US where maize (Zea mays L.) is grown in alleys between tree rows of either black walnut (Juglans nigra L.) or red oak (Quercus rubra L.). During a course of ten years, crop yields in rows adjacent to tree rows declined by 50% or more. With the experimental introduction of barriers to separate tree and crop root systems, yields in the rows near trees were equal to those of the center row (and monoculture). Irrespective of a high correlation between photosynthetically active radiation and net photosynthesis, shading did not have a major influence on crop yield. At this stage of system development (11 year old trees), influence of incident PAR on crop yield seems to be minimal. Subsequent papers in this series examine the sharing of belowground resources between trees and crops to quantify the competitive interactions that impact crop yields and their implications for economic return to the farmer.

Vegetation responses along edge-to-interior gradients in a high altitude tropical forest in peninsular India

A study was conducted in a high altitude tropical forest (shola forest) in peninsular India to examine the changes in floristic composition (mainly woody species regeneration) along an edge to interior gradient in relation to changes in edaphic and microenvironmental factors. Species inventory was taken in 25 m2 plots, established at 10 m intervals along edge to interior transects. The measured soil variables included pH, organic carbon, total nitrogen, available phosphorus, and soil moisture. Microenvironmental factors including light transmittance, relative humidity, and air and soil temperatures also were monitored. Both edaphic and microenvironmental factors exhibited strong patterns along the edge to interior gradient. Forest edges were characterized by higher light transmittance, higher air and soil temperatures, and lower relative humidity. Soil variables including pH, organic carbon, total nitrogen, available phosphorus, and moisture increased toward the forest interior. Significant increases in organic carbon (53.9%), total nitrogen (47%), and soil moisture (55%) indicated a relatively fertile forest interior compared with the forest edge. A definite floristic compositional pattern also was observed along the edge to interior gradient which was correlated to the edaphic and microenvironmental variables as revealed by canonical correspondence analysis. It appears that edge effects in these high altitude forests penetrate to a distance of 15–30 m. Further, edaphic factors have an important influence on woody species regeneration, perhaps much more than microenvironmental factors. This indicates that any disturbance that significantly exposes theforest floor, thus lowering soil moisture and altering soil nutrient status, can adversely affect the regeneration of many of the shola species.

Allelopathy in black walnut (Juglans nigra L.) alley cropping. I. Spatio-temporal variation in soil juglone in a black walnut-corn (Zea mays L.) alley cropping system in the midwestern USA.

A study was conducted to quantify the spatial and temporal variation in soil juglone (5-hydroxy-1,4-naphthoquinone) in a 10-year-old black walnut (Juglans nigra L.)–corn (Zea mays L.) alley cropping system. Two treatments (‘root barrier’ and ‘no barrier’) were applied to determine if soil juglone in the alley can be minimized by preventing black walnut root growth into the alley. Although no significant seasonal variation in soil juglone existed, a distinct spatial pattern was observed. Juglone concentration decreased as much as 80% as the distance increased to 4.25 m from the tree row. Installation of polyethylene root barriers minimized juglone concentration to trace levels in the alley. However, this treatment increased juglone levels within the tree row as compared to the ‘no barrier’ treatment, probably as a result of increased rooting density within a limited volume of soil.

Comparison of minirhizotron and soil core methods for quantifying root biomass in a temperate alley cropping system

A study was carried out in southern Indiana, USA with the objective of comparing soil core sampling and the minirhizotron technique in quantifying fine root biomass and root distribution patterns in an alley cropping system with black walnut (Juglans nigra L.), northern red oak (Quercus rubra L.) and maize (Zea mays L.). Spatial variation in tree rooting pattern was investigated prior to planting maize. Tree fine root biomass was quantified at distances of 0, 1.1, 2.3, 3.5, and 4.3 m where 0 m represents the tree row and 4.3 m represents the middle of the alley. Root samples were collected to a depth of 90 cm using a hydraulic auger. Maize rooting pattern was determined 65 days after planting to the same depth. Using plexiglass access tubes installed near the actual soil core locations and a minirhizotron camera root images were recorded on a VHS tape. These images were later analyzed using a raster based GIS software (ERDAS-IMAGINE). Regression analysis revealed significant relationships between root surface area measurements from minirhizotron observations and fine root biomass data from soil coring for all species. Predicted fine root biomass data were also in close agreement with actual fine root biomass for all species examined. Maize root biomass was slightly, but not significantly, underestimated by the minirhizotron technique in the top 30 cm soil layer. No significant underestimation or overestimation of root biomass in surface or deeper soil layers was observed for the tree species. The results indicate that minirhizotron can be used in quantifying fine root biomass if site and species specific predictive models can be developed.

Carbon, water and nutrient flux in Maya homegardens in the Yucatán peninsula of México

The Yucatán Peninsula of México has shallow soils and receives low amounts of precipitation, and has therefore low agricultural potential. Lacking large-scale irrigation from rivers and adequate rainfall, the indigenous Maya groups maintain agricultural productivity by adapting a variety of practices. Multistrata homegardens, one of their agricultural systems, have provided goods for trade, sale and personal consumption for many centuries. Nevertheless, an understanding of the controlling biological factors and interactions within these systems can lead to yield improvements. Photosynthetic rates, water use and litter production for a variety of species have been studied in these gardens to determine how Maya management impacts resource flow, productivity and diversity. When irrigated, diurnal photosynthetic rates nearly doubled for Manilkara zapota and water use increased two fold for M. zapota and Cordia dodecandra. Total litter production in traditional homegardens varied from 1,000 to 4,000 kg ha−1 yr−1 and ten arboreal species were found to contribute more than 33% of total litterfall biomass. Nutrient concentrations in the leaves of the predominant species were analyzed; Meliococcusbijugatus and Spondias purpurea were found to contribute the largest quantities of N, P and C. Our research indicates that species and structural diversity are critical to sustainability of homegardens, allowing efficient use and transfer of carbon, nutrients and water.

Defining competition vectors in a temperate alley cropping system in the midwestern USA. 4. The economic return of ecological knowledge

We tested the economic value of ecological knowledge in a midwestern USA alley-cropping system where row crops are planted in alleys between fine hardwood trees grown for veneer. Economic models were constructed to compare among agroforestry designs as well as to compare agroforestry with traditional forest plantation culture and row crop monoculture and rotational management. The general modeling approach was to quantify production inputs and outputs, estimate costs and revenues, simulate tree growth and crop productivity in agroforestry configurations, and estimate discounted cash flows. We incorporated scenarios that controlled both above- and below-ground competition through appropriate management as found in our previous research. This research showed the importance of below-ground competition in determining crop yields and the period of time that crop income could be expected from the agroforestry interplanting. Net present values and internal rates of return showed that agroforestry systems were generally more favorable investments than traditional agriculture and forestry. More importantly, the use of simple management techniques targeted at reducing below-ground competition allowed longer cultivation of row crops, greatly increasing returns to the landowner. Thus, the economic benefit of understanding the ecological interactions within agroforestry plantings dictates that accurate assessment of agroforestry alternatives will require the modeling of agroforestry as an integrated, interactive system.