Shade Species Research Articles

Transpiration of Arabica Coffee and Associated Shade Tree Species in Sub-optimal, Low-altitude Conditions of Costa Rica

Sap flows of coffee (Coffea arabica L. cv ‘Costa Rica 95’) and associated timber trees (Eucalyptus deglupta or Terminalia ivorensis) or leguminous tree (Erythrina poeppigiana) were measured simultaneously during 12 months in 4-year-old coffee agroforestry systems in sub-optimal ecological conditions of Costa Rica. In the wet period, coffee and shade tree transpiration followed the daily patterns of photosynthetic photon flux density (PPFD) and reference evapotranspiration (ETo) while their transpiration was restricted at higher air VPD values (>1.5 kPa) registered during the dry period. Coffee transpired more per unit leaf area in full sun than under shade, an indication of higher environmental coffee stress in non shaded conditions. Nonetheless, coffee daily water consumption per hectare was generally higher under shade than in full sun due higher vegetative growth of shade-grown coffee plants. Minimum and maximum daily transpiration were 0.74 and 4.08 mm for coffee, 0.35 and 1.06 mm for E. deglupta, 0.70 and 2.10 mm for T. ivorensis and 0.13 and 0.79 mm for E. poeppigiana. Estimation of the annual combined water transpiration by coffee and shade trees was 20–250% higher than that of coffee grown in full sun. Nevertheless, there was no evidence that water use by associated trees decreased soil water availability for coffee and hence limited coffee transpiration in the dry season due to its relatively short length (3 months) and the high annual rainfall (over 3100 mm). In the sub-optimal, low altitude conditions of this experiment, E. deglupta was the optimum shade species as it maintained a more constant shade level throughout the year and ensured a better protection to coffee underneath than T. ivorensis and E. poeppigiana which underwent a complete defoliation during the adverse meteorological conditions of the dry period.

Tradeoff between shade adaptation and mitigation of photoinhibition in leaves of Quercus mongolica and Acer mono acclimated to deep shade

We investigated susceptibility to photoinhibition in leaves acclimated to different light regimes in intermediately shade-tolerant Japanese oak (Quercus mongolica Fisch. ex Turcz. var. crispula (Blume) Ohashi) and shade-tolerant Japanese maple (Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara), to elucidate adaptability to gap formation in leaves differing in shade acclimation. We hypothesized that there is a tradeoff between shade adaptation and capacity to mitigate photoinhibition associated with leaf morphology. We simultaneously measured chlorophyll fluorescence and gas exchange in seedlings that had been grown in full sunlight (open), 10% of full sun (moderate shade) and 5% of full sun (deep shade). Shade-tolerant A. mono adapted to deep shade through changes in leaf morphology, lowering its leaf mass per area (LMA), but Q. mongolica showed little change in LMA between moderate and deep shade. Photochemical quenching (qP) did not differ between species in full sunlight and moderate shade; however, in deep shade, qP of Q. mongolica was higher than that of A. mono, suggesting that Q. mongolica grown in deep shade is less susceptible to photoinhibition at gap formation. This is consistent with the finding that chronic photoinhibition 3 days after the transfer to full sunlight, indicated by the decrease in maximum photochemical efficiency, Fv/Fm, at predawn, was less in deep-shade-grown Q. mongolica than in deep shade-grown A. mono. In deep shade, the electron transport rate (ETR) of Q. mongolica was higher than that of A. mono, whereas thermal energy dissipation through photosystem II antennae, indicated by non-photochemical quenching, was lower in Q. mongolica than in A. mono. In deep shade, the greater ETR capacity in Q. mongolica in association with higher LMA and higher leaf N content could contribute to maintaining high qP and mitigating photoinhibition. These results indicate that, by maintaining a high electron transport capacity even in deep shade, the gap-dependent and intermediate-shade-tolerant Q. mongolica trades improved shade adaptation for higher growth potential when a gap event occurs.

Allometry, adult stature and regeneration requirement of 65 tree species on Barro Colorado Island, Panama

This study provides a community-level analysis of how regeneration requirement and adult stature are related to tree allometry (diameter, height and crown size) throughout post-seedling ontogeny on Barro Colorado Island, Panama. Comparing 65 species, gap species are taller, have higher diameter growth rates and occupy more low-canopy sites (≤10 m canopy height) than shade species at small diameters (≤10 cm dbh). For trees >10 cm dbh, diameter-height relationships and growth rates no longer differ between gap and shade species, but shade species have larger, particularly deeper, crowns than gap species. Species with tall adult stature have more slender stems with larger crowns compared with treelet and mid-canopy species starting at 5 cm dbh. From 10 to 40 cm dbh, diameter growth rate is also significantly greater for tall species. The consistent allometric differences between functional groups on a community level will, in part, determine vertical and horizontal stand structure.

Foraging ecology of howler monkeys in a cacao (Theobroma cacao) plantation in Comalcalco, Mexico

Recent evidence indicates that primate populations may persist in neotropical fragmented landscapes by using arboreal agroecosystems, which may provide temporary habitats, increased areas of vegetation, and connectivity, among other benefits. However, limited data are available on how primates are able to sustain themselves in such manmade habitats. We report the results of a 9-month-long investigation of the feeding ecology of a troop of howler monkeys (n = 24) that have lived for the past 25 years in a 12-ha cacao plantation in the lowlands of Tabasco, Mexico. A vegetation census indicated the presence of 630 trees (> or =20 cm diameter at breast height (DBH)) of 32 shade species in the plantation. The howlers used 16 plant species (13 of which were trees) as sources of leaves, fruits, and flowers. Five shade tree species (Ficus cotinifolia, Pithecellobium saman, Gliricidia sepium, F. obtusifolia, and Ficus sp.) accounted for slightly over 80% of the total feeding time and 78% of the total number trees (n = 139) used by the howlers, and were consistently used by the howlers from month to month. The howlers spent an average of 51% of their monthly feeding time exploiting young leaves, 29% exploiting mature fruit, and 20% exploiting flowers and other plant items. Monthly consumption of young leaves varied from 23% to 67%, and monthly consumption of ripe fruit varied from 12% to 64%. Differences in the protein-to-fiber ratio of young vs. mature leaves influenced diet selection by the monkeys. The howlers used 8.3 ha of the plantation area, and on average traveled 388 m per day in each month. The howlers preferred tree species whose contribution to the total tree biomass and density was above average for the shade-tree population in the plantation. Given the right conditions of management and protection, shaded arboreal plantations in fragmented landscapes can sustain segments of howler monkey populations for many decades.

Growth Response of Three Turfgrass Species to Nitrogen and Trinexapac-ethyl in Shade

Previous research on the potential of the gibberellin inhibiting growth regulator trinexapac-ethyl (TE) [4-(cyclopropyl-α-hydroxy-methylene)-3,5-dioxocyclohexanecarboxylic acid ethyl ester] to improve quality and density of shaded turfgrass has been conducted under neutral-density shade. However, some phytochrome-mediated growth responses of turfgrass, such as tillering, are different under deciduous shade versus neutral-density shade. The objectives of this study were to investigate 1) whether TE would result in improved stand density and quality of turfgrass grown under deciduous shade as has been observed under neutral-density shade and 2) the shade tolerance of sheep fescue (Festuca ovina L. `Quatro') compared to tall fescue (Festuca arundinacea Schreb. `Plantation'), and rough bluegrass (Poa trivialis L.). Trinexapac-ethyl at either 0 or 29 kg·ha–1 a.i. and nitrogen at 12 or 36 kg·ha–1 were applied on 23 May, 3 July, and 15 Aug. 2003 and 21 May 2004 to each species in a randomized complete block design under deciduous shade (about 9% of full sun). Clipping yield, color, and density data were collected for 6 weeks after the May applications in each year. Visual quality was assessed for 6 weeks after application in 2004 only. In 2003, TE significantly reduced clipping yields by 35% to 50% on sheep fescue, 58% to 76% on tall fescue and 55% to 80% on rough bluegrass. However, in 2004, yield reduction was 0% to 50% for all three species and there was no interaction between week, TE, and species. `Plantation' tall fescue had the highest overall visual quality and density. Sheep fescue also provided an acceptable quality turf stand. TE application did not significantly impact the quality of these species. Rough bluegrass performance was unacceptable, and high rate applications of TE to this species in shade resulted in significant (P < 0.05) losses in density. Trinexapac-ethyl application, based on the results of this study, may not enhance turf quality of cool season grasses grown under dense tree shade.

Fertilizer Applications on Establishment and Growth of Three Groundcover Species in Sun and Shade

Abstract Use of groundcovers in the landscape is often limited by slow establishment when compared to turf. Hedera helix L. (English ivy), Euonymus fortunei ‘Coloratus’ (Turcz.) Hand.-Mazz. (purpleleaf wintercreeper euonymus) and Liriope spicata Lour. (creeping lily-turf) were evaluated in full sun and 60% shade to determine the effects of fertilizer applications on establishment and growth. Limited fertilizer × species interaction was detected, which may have been due, in part, to the soil incorporation of mushroom compost. Establishment and survivability of Hedera in full sun may have been influenced by lack of acclimation from shade to sun. There was no significant difference in establishment rates between Liriope and Euonymus; however, under shade Euonymus did not develop its characteristic fall color. Hedera was established in one season under 60% shade and, of these three species, can be considered the most competitive with turf under similar conditions for low traffic areas of the landscape.

Response of mitochondria to light intensity in the leaves of sun and shade species

ABSTRACTThe present authors have shown previously that both respiration rates and in vivo activities of the alternative oxidase (AOX) of leaves of Alocasia odora, a shade species, are lower than those in sun species, thereby optimizing energy production under limited light conditions (Noguchi et al., Australian Journal of Plant Physiology 28, 27–35, 2001). In the present study, mitochondria isolated from A. odora leaves were examined in order to investigate the biochemical basis for the differences in respiratory parameters. Alocasia odora and spinach plants were cultivated under both high and low light intensities, mitochondria were isolated from their leaves, and their respiratory properties compared. Mitochondrial content of leaf extracts from the two species was estimated using fumarase activities and antibody detection of porin (the voltage‐dependent anion channel of the outer mitochondrial membrane). On a mitochondrial protein basis, spinach leaves showed higher capacities of the cytochrome pathway and cytochrome c oxidase (COX) than A. odora leaves. However, on a mitochondrial protein basis, A. odora showed higher capacities of AOX, which had a high affinity for ubiquinone when activated by pyruvate. Alocasia odora also had larger amounts of mitochondrial protein per leaf dry weight, even under severely shaded conditions, than spinach. Lower growth light intensity led to lower activities of most pathways and proteins tested in both species, especially glycine‐dependent oxygen uptake. In the low light environment, most of the AOX protein in A. odora leaves was in its inactive, oxidized dimer form, but was converted to its reduced active form when plants were grown under high light. This shift may prevent over‐reduction of the respiratory chain under photo‐oxidative conditions.

Hydraulic efficiency of the leaf venation system in sun- and shade-adapted species.

We tested the hypothesis that leaf hydraulics is correlated with the light adaptation of different plant species and specifically that the hydraulic resistance of the leaf venation (Rvenation) is lower in sun- than in shade-adapted species. Rvenation was measured in six sun- and six shade-adapted species with a high-pressure flow meter (HPFM). The number of conduits at the proximal third of the midrib was counted and the diameter of the widest conduits together with vein density were measured. Rvenation was higher in shade species than in sun species and it was negatively correlated with the mean diameter of the widest conduits. Maximum leaf conductance to water vapour recorded for the different species was negatively correlated with the corresponding Rvenation. Sun-adapted species coping with the high water demand typical of sunny habitats appeared to have developed a highly efficient conducting system to supply living mesophyll cells with water. In contrast, species adapted to shady habitats showed higher Rvenation values according to their lower need for investment of carbon into producing wide conduits in the leaf.

Butterfly community composition and conservation in and around a primary woodland of Mount Fuji, central Japan

A study was conducted in and around the Aokigahara primary woodland of Mount Fuji, central Japan, to clarify butterfly community structure along the environmental gradients of human disturbance, shade, and plant species richness, with a view to formulating conservation strategies for the community. The composition and abundance of butterfly species were recorded during 1999 along transects in three habitats: woodland, woodland edge and open land. Two sites were selected for each habitat. Canonical correspondence analysis (CCA) showed that community structure was correlated with both disturbance and shade. However, neither relationship was significant when disturbance and shade were analyzed as covariables (partial CCA), suggesting that the combined effect of the two variables played an important role in determining community structure. In contrast, CCA showed no significant relationships between butterfly community structure and plant or hostplant species richness. Correlation analyses between species characteristics and the CCA scores showed that species occurring in more disturbed habitats were more voltine, had a wider hostplant range and were associated with lower shade tolerance than those occurring in less disturbed habitats, and species occurring in darker habitats with higher shade tolerance. Generally, the structure and characteristics of the butterfly community could be understood well in terms of the habitat templet theory. To conserve endangered species on the Red List of Japan, and to maintain butterfly diversity in and around the woodland, this study suggests a need not only to conserve the primary woodland area, but most importantly to maintain the surrounding forest edge including areas of semi-natural grassland.

Using environmental and growth characteristics of plants to detect long-term changes in response to atmospheric pollution: some examples from British beechwoods

This study uses the Ellenberg system of plant indicator values, along with Grime et al.'s plant growth strategy values, to investigate the nature of temporal changes in the composition of ground flora in two beechwoods in the Cotswolds region of the UK, currently receiving atmospheric inputs of nitrogen in excess of critical loads. The woods, first surveyed in the early 1960s, were resurveyed in 1998 using the original sampling protocol. Temporal changes in the abundance of individual species at Blackstable West Wood indicate changes in light over time, although decreases in sun species, and both increases and decreases in shade species suggest that this change has been patchy. Analysis of changes in plant community as represented by weighted and unweighted quadrat Ellenberg and CSR scores have yielded more significant results. Blackstable West Wood shows increases in nitrophilic, moist-soil and competitive species accompanied by a decline in stress-tolerant species. In Buckholt Top Wood there has been an increase in sun and moist-soil species, a decrease in competitive species and, when weighted Ellenberg scores are considered, an increase in acid-tolerant species. These changes indicate both the impact of woodland management by selective felling and an underlying influence of enhanced atmospheric deposition especially of nitrogen pollutants. It is concluded that quadrat mean scores are a useful tool especially where few individual species have undergone large temporal changes in abundance. However, lack of correlations between quadrat Ellenberg scores for pH and light on one hand, and their equivalents measured in the field, i.e. soil pH and surrogates for light, such as distance to the nearest tree and tree density, suggest that this approach may not be sensitive enough to pick up small-scale, within site variations. Although harder to interpret, plant strategy scores were found to be a useful additional descriptor, encapsulating a plant's response to a range of environmental factors.

Patterns of leaf surface wetness in some important medicinal and aromatic plants of Western Himalaya

The present study aims at investigating the role of leaf morphological features and their relation to leaf surface wettability for important medicinal and aromatic plants of Western Himalaya. The surface features related to leaf wettability were studied in 30 plant species representing 21 different families growing under open and shade conditions. The leaf surfaces with the highest density of trichomes and stomata per unit area were found to be the least wettable, regardless of condition type. Most of the species of both conditions were hypostomatic, and per unit area concentration of stomata contributes more than stomatal size to stomatal area index. Leaf surfaces of open condition species were more water repellent with higher stomatal density, and had lower water droplet retention than shade species. It is suggested that leaf morphological features (stomata and trichome) had a strong influence in reducing the leaf area with surface moisture, which could be correlated with the frequency and duration of leaf wettability in a given condition.

Effects of short-term irradiation on photoinhibition and accumulation of mycosporine-like amino acids in sun and shade species of the red algal genus Porphyra

The effect of irradiance (40 and 840 μmol photons m −2 s −1) of short-term (48 h) irradiation on photosynthetic activity (estimated as oxygen evolution and as chlorophyll fluorescence), specific absorption and fluorescence excitation spectra, photosynthetic pigment accumulation (chlorophyll a and biliproteins) and UV-absorbing compounds (mycosporine-like amino acids, MAAs) was investigated in sun and shade species of the red algal genus Porphyra collected in Trondheimsfjord (Norway). In the sun type, high irradiance exposure (840 μmol photons m −2 s −1) did not alter the Chl a concentration, however, exposure to a lower irradiance (40 μmol photons m −2 s −1) for 48 h significantly increased the chlorophyll concentration. The content of MAAs was significantly higher in the suntype than in the shade type algae. Porphyra-334 is the main MAA in this species followed by shinorine. The total content of MAAs significantly ( P<0.05) increased in the sun type after 48 h exposure to both high and low irradiances. However, in the shade type, porphyra-334 significantly decreased ( P<0.05) after both high and low irradiance exposure. Photosynthetic activity (as oxygen evolution) and the optimal quantum yield ( F v/ F m), as an indicator of photoinhibition, decreased under low and high irradiance in the shade type algae and no full recovery was observed when the algae were transferred to very low irradiation.The sun type algae presented a higher capacity of acclimation to increased irradiance than the shade type algae. This high acclimation of sun type algae to short term high irradiance exposure (48 h) is explained by the higher thermal dissipation. This was estimated as the ratio of nonphotochemical quenching related to the light dose ( q N:dose) and by the accumulation of MAAs.

Short-term light and leaf photosynthetic dynamics affect estimates of daily understory photosynthesis in four tree species.

Instantaneous measurements of photosynthesis are often implicitly or explicitly scaled to longer time frames to provide an understanding of plant performance in a given environment. For plants growing in a forest understory, results from photosynthetic light response curves in conjunction with diurnal light data are frequently extrapolated to daily photosynthesis (A(day)), ignoring dynamic photosynthetic responses to light. In this study, we evaluated the importance of two factors on A(day) estimates: dynamic physiological responses to photosynthetic photon flux density (PPFD); and time-resolution of the PPFD data used for modeling. We used a dynamic photosynthesis model to investigate how these factors interact with species-specific photosynthetic traits, forest type, and sky conditions to affect the accuracy of A(day) predictions. Increasing time-averaging of PPFD significantly increased the relative overestimation of A(day) similarly for all study species because of the nonlinear response of photosynthesis to PPFD (15% with 5-min PPFD means). Depending on the light environment characteristics and species-specific dynamic responses to PPFD, understory tree A(day) can be overestimated by 6-42% for the study species by ignoring these dynamics. Although these overestimates decrease under cloudy conditions where direct sunlight and consequently understory sunfleck radiation is reduced, they are still significant. Within a species, overestimation of A(day) as a result of ignoring dynamic responses was highly dependent on daily sunfleck PPFD and the frequency and irradiance of sunflecks. Overall, large overestimates of A(day) in understory trees may cause misleading inferences concerning species growth and competition in forest understories with < 2% full sunlight. We conclude that comparisons of A(day) among co-occurring understory species in deep shade will be enhanced by consideration of sunflecks by using high-resolution PPFD data and understanding the physiological responses to sunfleck variation.

Antioxidant capacities of green and cyanic leaves in the sun species, Quintinia serrata.

Anthocyanins scavenge harmful reactive oxygen species (ROS), and may provide antioxidative protection within plant cells. For the shade species Elatostema rugosum A. Cunn., it was recently established that red-leaved morphs held a significant antioxidant advantage over green morphs. It is not known whether anthocyanins confer the same protection in sun species that are exposed to elevated levels of irradiation and thus greater photooxidative stress. Quintinia serrata A. Cunn., a canopy tree species, is polymorphic for anthocyanin expression. Levels of antioxidants in leaves of red and green morphs of Q. serrata were identified, and activities quantified using enzymatic and α,α-diphenyl-β-picrylhydrazyl (DPPH) assays and cyclic voltammetry. Both morphs contained similar levels of superoxide dismutase, catalase, ascorbic peroxidase and hydroxycinnamic acids, and showed similar ranges in DPPH scavenging activity and voltammetric profiles. Anthocyanin concentrations increased with leaf age in red morphs and were positively correlated with the extent of DPPH scavenging. However, hydroxycinnamic acids were the most active methanol-extractable fraction in both morphs, contributing 27% more to the overall low-molecular-weight antioxidant pool in the green morph than in the red. We conclude that anthocyanin biosynthesis can enhance, but is not a prerequisite for, oxidative protection in Q. serrata leaves. The possibility that anthocyanins provide a versatile mechanism for photoprotection is discussed.

Why do species of woody seedlings change rank in relative growth rate between low and high irradiance?

It is often assumed that if a plant species has a higher relative growth rate (RGR) than another species in deep shade, it will have a lower RGR at high irradiance (Spurr & Barnes 1980; Thomas & Bazzaz 1999; Walter 1973). In other words, species change rank (crossover) between low and high irradiance. This idea was suggested chiefly by the finding that in deep shade the massbased net photosynthetic rates of the leaves of shade plants exceed those of sun plants, while at high irradiance the reverse is true (Bjdrkman & Holmgren 1963; Boardman 1977; Givnish 1988). This finding has been assumed to scale up to the level of whole-plant RGR (Shugart 1984). In the past decade, however, a contrary view concerning RGRs has emerged, the idea that if a plant grows faster than another at high irradiance it will also do so in the shade (Kitajima 1994; Poorter 1999). According to this later view, light plays a role in maintaining the mixture of forest species only through the well established trade-off between survival rate in deep shade and RGR in bright light (Kitajima 1994, 1996). Similar experimental studies of RGR responses to irradiance for woody seedlings report surprisingly different results. Here we indicate why such disparate results have been produced. We provide a simple analytical approach to understanding why crossovers should occur among particular species at particular stages of ontogeny. This approach is useful for understanding the maintenance of forest species richness, as well as for interpreting plant specialization in physiology and morphology to contrasting irradiance regimes.

Costs of protein turnover and carbohydrate export in leaves of sun and shade species

In mature leaves, adenosine triphosphate (ATP) provided by respiration is used for maintenance of leaves and carbohydrate export (starch mobilisation and phloem loading). The main maintenance processes of mature leaves include turnover of existing structures (e.g. proteins and membranes) and the trans-membrane transport of solutes. In order to mechanistically estimate these costs in mature leaves and compare them between sun and shade species, we measured the costs of protein turnover and carbohydrate export in mature leaves of Alocasia odora (Lodd.) Spach., a shade-tolerant species, and Phaseolus vulgaris L., a sun species, in the night. We estimated the rate of ATP production from the rate of O 2 uptake, taking account of the contribution of the alternative respiratory pathway, assessed by the O 2 isotope fractionation technique. The energy consumption of protein turnover was estimated from the decrease in the rate of ATP production in the presence of an inhibitor of cytosolic protein synthesis. Examination of the anatomy of the minor veins in the leaves revealed that A. odora employs symplastic phloem loading, while P. vulgaris was reported to employ apoplastic loading. Based on these phloem loading types and the difference between the rate of carbohydrate decrease and the CO 2 efflux rate in the leaves, we calculated the ATP cost for carbohydrate export. The costs estimated for two processes amounted to about 40% of the ATP production rate in A. odora and 80% in P. vulgaris. The absolute costs for the two processes in the leaves of A. odora were much lower than those of P. vulgaris. Both the cost of protein turnover per unit leaf nitrogen and that of carbohydrate export per exported carbon were lower in the leaves of A. odora. Low ATP consumption rates by these cellular processes would explain the low respiratory rate in A. odora.

Activities of the cyanide-resistant respiratory pathway in leaves of sun and shade species

Higher plant mitochondria have a cyanide-resistant alternative respiratory pathway of electron transport (AP) that is not coupled to proton translocation. To characterise the ecophysiology of this apparently ‘wasteful’ pathway, we constructed a system consisting of a gas-phase oxygen electrode and an air sampling line for measurement of stable oxygen isotope ratios. With this system, we were able to measure respiratory rates of a small amount of leaf segments of ca 0.6 g fresh weight, and collect about 100 L of the air from the oxygen electrode chamber several times. The 18 O/16 O ratio in the air samples was measured by mass spectrometry. The activity of AP was estimated based on the isotopic discrimination of 18 O. We used the leaves of Alocasia odora (Lodd.) Spach., a shade species, and Spinacia oleracea L. and Phaseolus vulgaris L., sun species. These plants were grown at two levels of photosynthetically active photon flux density (PPFD). Three main findings were: (1) in the leaves of A. odora, the contribution of AP was less than 10% of the total respiratory rate, irrespective of growth PPFD; (2) for the sun species grown at high PPFD, the contribution of AP in the leaves was about 40% early in the night, but decreased dramatically late in the night; and (3) when S. oleracea was grown at low PPFD, the contribution of AP in the leaves declined. The low activity of AP in the leaves of A. odora suggests that the efficiency of adenosine triphosphate (ATP) production (ATP/O 2 ) of this species is high. This may be especially important in shaded environments where input of light energy is low. We also suggest that, in the leaves of sun species, ATP/O 2 changes depending on the conditions.

Research methods for multistrata agroforestry systems with coffee and cacao: recommendations from two decades of research at CATIE

This paper reviews the research themes and methodologies used by CATIE in agroforestry research with shade trees over coffee (Coffea arabica) and cacao (Theobroma cacao) during the past 20 years. Initially research focused on characterization and production studies (of crop and timber including border areas) of traditional systems using temporary and permanent sample plots on private farms. The assessment area of traditional shade-coffee (or cacao) systems should be the whole plot, including the border areas, and not some subjectively selected central area which supposedly represents unit area productivity. Uncontrolled crop, tree, and management heterogeneity limited extrapolation of early on-farm research results to other farmers' fields. Replicated case studies of best bet technologies (traditional or experimental) on different farms are often preferable to the use of formal experimental designs. On-station research included the use of systematic spacing designs to test extreme shade tree density treatments of coffee. Most nutrient cycling studies were also carried out on-station, using service and timber shade species over coffee and cacao to evaluate the ability of these agroforestry systems to maintain nutrient reserves and diversify production. Plot size (even 36 × 36 m) was limiting for long term research because of inter-plot interference, both below- and above ground, when using fast growing, tall timber trees as shade. These experiences suggest a minimum plot size of 2,500 m2. Individual tree designs and tree-crop interface studies (e.g. regression analysis of data taken along transects) are promising experimental/sampling approaches that need further development. The principal research thrusts proposed for the next five years are bio-physical process research on coffee responses to shade and competition with trees (growth, carbon allocation, phenology, disease-pest tolerance, yields and coffee quality effects) and socioeconomic analyses of both traditional and new or improved shade – coffee combinations vs. monocultures.

Monitoring of oxygenic and anoxygenic photosynthesis in a unicyanobacterial biofilm, grown in benthic gradient chamber

In order to assess the role of cyanobacteria in the formation and dynamics of microenvironments in microbial mats, we studied an experimental biofilm of a benthic, halotolerant strain, belonging to the Halothece cluster of cyanobacteria. The 12-week-old biofilm developed in a sand core incubated in a benthic gradient chamber under opposing oxygen and sulfide vertical concentration gradients. At the biofilm surface, and as a response to high light irradiances, specific accumulation of myxoxanthophyll was detected in the cells, consistent with the typical vertical distribution of sun versus shade species in nature. The oxygen turn-over in terms of gross photosynthesis and net productivity rates was comparable to oxygen dynamics in natural microbial mats. Sulfide blocked O 2 production at low irradiances in deep biofilm layers but the dynamics of H 2S and pH demonstrated that sulfide removal by anoxygenic photosynthesis was taking place. At higher irradiances, as soon as H 2S was depleted, the cells switched to oxygenic photosynthesis as has been postulated for natural communities. The similarities between this experimental biofilm and natural benthic microbial mats demonstrate the central role of cyanobacteria in shaping microenvironmental gradients and processes in other complex microbial communities.

Monitoring of oxygenic and anoxygenic photosynthesis in a unicyanobacterial biofilm, grown in benthic gradient chamber.

In order to assess the role of cyanobacteria in the formation and dynamics of microenvironments in microbial mats, we studied an experimental biofilm of a benthic, halotolerant strain, belonging to the Halothece cluster of cyanobacteria. The 12-week-old biofilm developed in a sand core incubated in a benthic gradient chamber under opposing oxygen and sulfide vertical concentration gradients. At the biofilm surface, and as a response to high light irradiances, specific accumulation of myxoxanthophyll was detected in the cells, consistent with the typical vertical distribution of sun versus shade species in nature. The oxygen turn-over in terms of gross photosynthesis and net productivity rates was comparable to oxygen dynamics in natural microbial mats. Sulfide blocked O(2) production at low irradiances in deep biofilm layers but the dynamics of H(2)S and pH demonstrated that sulfide removal by anoxygenic photosynthesis was taking place. At higher irradiances, as soon as H(2)S was depleted, the cells switched to oxygenic photosynthesis as has been postulated for natural communities. The similarities between this experimental biofilm and natural benthic microbial mats demonstrate the central role of cyanobacteria in shaping microenvironmental gradients and processes in other complex microbial communities.