The roles of abscisic acid and ethylene in the abscission and senescence of cocoa flowers

Cocoa flowers have a limited period of longevity; more than 90% of unpollinated flowers abscised within 32 h after anthesis. Abscisic acid (ABA) levels increased significantly prior to abscission. By 21 h after anthesis, ABA levels had increased almost 10-fold, and by 32 h flowers had 20-fold higher levels of ABA than at anthesis. Fluridone completely inhibited both the increase in ABA, the formation of an abscission zone, and the abscission and senescence of flowers. In contrast, ethylene production increased only slightly 21 h after anthesis and was only 2-fold higher after 32 h. Aminoethoxyvinylglycine (AVG) delayed but did not prevent abscission. In cocoa flowers, ABA is the primary regulator of abscission; ethylene accelerates abscission but only in the presence of ABA. Naphthalene acetic acid (NAA) treatment of flowers at anthesis prevented abscission zone formation and flower abscission, but did not induce fruit set. All parts of the NAA-treated flower except the pedicel senesced after 6 days. NAA+AVG treatment only delayed, whereas fluridone treatment completely prevented flower senescence.     

Effects of an experimental drought on the functioning of a cacao agroforestry system,Sulawesi, Indonesia

Agroforestry systems may play a critical role in reducing the vulnerability of farmers' livelihood to droughts as tree‐based systems provide several mechanisms that can mitigate the impacts from extreme weather events. Here, we use a replicated throughfall reduction experiment to study the drought response of a cacao/Gliricidia stand over a 13‐month period. Soil water content was successfully reduced down to a soil depth of at least 2.5 m. Contrary to our expectations we measured only relatively small nonsignificant changes in cacao (?11%) and Gliricidia (?12%) sap flux densities, cacao leaf litterfall (+8%), Gliricidia leaf litterfall (?2%), soil carbon dioxide efflux (?14%), and cacao yield (?10%) during roof closure. However, cacao bean yield in roof plots was substantially lower (?45%) compared with control plots during the main harvest following the period when soil water content was lowest. This indicates that cacao bean yield was more sensitive to drought than other ecosystem functions. We found evidence in this agroforest that there is complementary use of soil water resources through vertical partitioning of water uptake between cacao and Gliricidia. This, in combination with acclimation may have helped cacao trees to cope with the induced drought. Cacao agroforests may thus play an important role as a drought‐tolerant land use in those (sub‐) tropical regions where the frequency and severity of droughts is projected to increase.     

Conservation Value of Cacao Agroforestry Systems for Terrestrial Herbaceous Species in Central Sulawesi,Indonesia

Tropical secondary forest and agroforestry systems have been identified as important refuges for the local species diversity of birds and other animal groups, but little is known about the importance of these systems for terrestrial herbs. In particular, few studies report how the conversion from tropical forest to technified cacao plantation affects the species richness and the community structure of herbs. We conducted surveys in 43 cacao plantations along the border of the Lore Lindu National Park in Central Sulawesi, ranging from agroforests to technified cacao, categorizing the plantations as rustic cacao, planted shade cacao, and technified cacao. We recorded 91 herb species. Of the 74 species determined to species level, 21 were also found in natural forests, while 53 were recorded only in agricultural habitats. Araceae was the most forest‐dependent plant family while Asteraceae included the highest number of nonforest species. Overall, the presence of forest species was confined to moderately intensively managed rustic and planted shaded plantations. Distance from the forest, which has been identified as a crucial parameter for the diversity and composition of other taxa in cacao agroforests, only played a minimal role for herbs. Our study suggests that native forest herbs maybe more vulnerable to forest conversion than animal groups. The intensification of cacao plantation management increases the presence of weedy species to the detriment of native forest species.     

Effect of Crinipellis perniciosa Infection on Abscission of Cacao Cotyledons, Reserve Mobilization and Dry Matter Partitioning

Infection by Crinipellis perniciosa on cacao seedlings prevented the abscission of cotyledons which normally abscise within 32–61 days from seed sowing. The early growth and development of control and artificially inoculated seedlings was investigated for root dry mass (RDM), shoot dry mass (SDM), shoot‐to‐root ratio (SRR), total dry mass (TDM), cotyledonary dry mass (CDM) and cotyledonary water content (CWC). Cotyledonary reserve mobilization was estimated at 69 and 56% at 5 weeks and 79 and 64% at 8 weeks in control and infected seedlings, respectively. Infection induced the enlargement of tissues but was not accompanied by an increase in dry mass. Infection both delayed and reduced the utilization of cotyledonary food reserves resulting in slower accumulation of RDM, SDM and TDM culminating in a reduction of growth by a factor of 1.6 at 8 weeks. However, the SRR of inoculated plants was similar to that of control plants, suggesting that the balance between the root and shoot systems exists which is unchanged by the effect of the pathogen. CWC was decreased upon senescence to an average of 65% at abscission. The prevention of cotyledonary abscission in infected seedlings is suggested to be due in part to the CDM remaining above a critical level of 20% of the initial CDM. The implication of this response in relation to the infection biology of the pathogen is discussed.     

Phenol content, acidic peroxidase and IAA-oxidase during somatic embryogenesis in Theobroma cacao L.

Calli were induced in cacao cotyledon explants on a half-strength Murashige and Skoog medium containing 6 × 10-2 g m-3 saccharose and various combinations of 2,4-dichlorophenoxyacetic acid (2,4-D) with kinetin (kin), benzylaminopurine (BAP) or 2-isopentenylphosphate (2-iP). Experiments were carried out on two clones of cacao differing in their susceptibility to black pod disease. The highest percentage of explants forming callus and the most rapid callus development were obtained with 10-6 g m-3 2,4-D and 0.5× 10-6 g m-3 kin. Somatic embryogenesis and rhizogenesis were induced by transferring 3-week-old callus in a half strength Murashige and Skoog medium containing 3 × 10-2 g m-3 saccharose and NAA or IBA in the 0 to 5 × 10-6 g m-3 concentration range. No differentiation could be observed when the medium was supplemented with kin or BAP. The conversion of callus into somatic embryos and roots was accompanied by a drop in phenol content and an increase in peroxidase and IAA-oxidase activities. Moreover, cell differentiation was characterized by the persistence in the callus of one acidic soluble isoperoxidase which was not detected in nondifferentiating callus. Although some differences were noticed between the clones, alterations responsible for cell differentiation were the same in both genotypes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Neighbor Trees: Shade, Intercropping, and Cacao in Ecuador

Previous studies have shown that shade trees in cacao and coffee are important habitats for inter-American migratory birds. A survey of 21 cacao farmers along the northern, central, and southern Pacific Coast of Ecuador found that shade trees are associated with unirrigated, traditional cacao varieties and low levels of chemical inputs. Farmers stressed the importance of shade for managing soil moisture and soil fertility, and for managing some weeds and diseases. Most of the shade trees were not wild forest trees, but had been planted and protected by the farm families. Many other trees are intercropped with cacao for economic reasons, not related to shade. Chocolate manufacturers, consumers, and environmental activists can encourage farmers to maintain shade canopies by paying a premium for the traditional, shade-loving, high-quality aromatic cocoa varieties.     

Effects of plant water deficit on the daily carbon balance of leaves of cacao seedlings

The daily carbon balance of individual source leaves of Theobroma cacao L. seedlings was measured at 2- to 3-day intervals during a 19-day period of increasing plant water deficit and during an 8-day period of recovery following rewatering. In each case, responses of stressed seedlings were compared to those of irrigated controls. Leaves of irrigated cacao seedlings assimilated approximately 41 mg carbohydrate dm^-2 during 12-h photoperiods, and exported an average of 34 mg carbohydrate dm^-2 during 24-h measurement cycles. The rate of carbon export from cacao leaves was sharply reduced as leaf water potential (ψ) declined between -0.8 and -2.0 MPa. Further, the rate of export was closely associated with the net assimilation rate (A), with export capacity being severely reduced as A fell to near zero. Net accumulation of dry matter occurred as long as A remained greater than approximately 20 mg carbohydrate dm^-2 over the 12-h photoperiod, but at lower assimilation rates, export exceeded concomitant assimilation. Carbon export continued at the expense of leaf carbon reserves as photoassimilation fell to near zero during periods of severe water stress (ψ < -2.0 MPa). Night respiration rate was independent of plant water status.     

Chlorophyll fluorescence parameters characterizing the development of two cacao genotypes infected by witches' broom

Greenhouse-grown susceptible 20-d-old seedlings of Theobroma cacao genotypes Catongo and tolerant genotype SCA6xCatongo were inoculated with a mixture of isolates of Crinipellis perniciosa, the causal agent of witches' broom. The characteristics of chlorophyll a fluorescence emission were monitored during leaf ontogeny using a portable system PAM-2000. In both inoculated and non-inoculated genotypes, significant differences were found for the effective quantum yield values of photosystem (PS) 2 (F/Fm') at the B (7 to 14-d-old), D (21 to 30-d-old), and E (>30-d-old) stages of leaf development, and in quantum yield of the non-cyclic photosynthetic electron transport between PS2 and PS1 [qp(Fv/Fm)] and quencher efficiency [(Fm-Ft)/F0] at the B, C (15 to 20-d-old) and D stages. Intergenotypic differences were found only for the [qp(Fv/Fm)] and [(Fm-Ft)/F0] values at the E stage, and for fluorescence quenching (Fm-Ft) at the B and E stages. Highly significant inter- and intragenotype relationships were found between the rate of photosynthetic electron transport to PS2 (Amax) and maximum fluorescence during actinic irradiation (Fm'). Also, each of the highly significant relationships between (Fm-Ft) and Amax, [(Fm-Ft)/F0] and F/Fm', and between [(Fm-Ft)/F0] and Amax were represented by a general model, independent of treatments. Therefore, alterations in energy distribution in the radiant energy collector complex interior of PS2 and reduction in absorption of photosynthetically active radiation were observed in the infected plants, mainly in the hybrid at the C stage. Also, variations were found in the noncyclic photosynthetic electron transport at the B and C stages in the infected Catongo.     

Heritability of Phenols in the Resistance of Theobroma cacao against Phytophthora megakarya, the Causal Agent of Black Pod Disease

The black pod disease caused by Phytophthora megakarya is responsible for 80% of the cocoa production loss in Cameroon. To assess the resistance of cocoa plants against this pathogen, necrotic lesions, phenolic content and qualitative alteration of phenolics were conducted in ICS84 and ICS95 clones (two Trinitario introduced from Trinidad) and their hybrids (families F30 and F25) derived from reciprocal cross breeding between these two parental clones after inoculation. The existence of strong hybrid vigour has been shown. Ninety percentage of the hybrid's genotypes manifested a positive heterosis effect for the development of lesion size. This suggests the existence of hybrid vigour with a genetic additive effect. F3086, F2509, F2552 and F2586 hybrids were characterized by localized lesions. Those hybrids genotypes can be considered as elite clones. In relation to analysis of total phenolics and lesion size, no maternal effect was detected in the transmission of these characters. A significant and negative correlation (r = −0.683) (P < 0.01) has been observed between necrosis evolution and phenolics accumulation. The values of the heritability of lesion size and the total phenolic content in offsprings did not permit to show the maternal effect. Qualitative analyses of phenolics showed high flavonones content in cocoa leaves. Qualitative analyses of phenolics in ICS84, ICS95 clones and their hybrids showed a modification of the phenolics profiles, notably concerning apigenin and luteolin derivatives due to the inoculation. These compounds, along with others that were not identified, could have a role in the reaction and mechanism of defence of cocoa against P. megakarya.     

浙江天童常绿植物当年生与往年生叶片性状的变异与关联

植物叶片性状随叶龄的变化是植物生活史策略的体现, 反映了植物叶片的物质投资和分配方式。该研究通过在个体和物种2个水平, 比较浙江天童1 hm ²样地内常绿阔叶树种的平均叶面积(MLA)、比叶面积(SLA)和叶片干物质含量(LDMC)在当年生和往年生叶片间的差异和关联, 探究叶片物质分配策略在异龄叶间的变化, 并分析叶龄对植物叶片性状, 特别是叶片面积建成消耗的影响。结果显示: 1)在个体和物种水平上, MLA变异系数最大(个体: 79.5%; 物种: 66.5%), SLA次之(个体: 28.1%; 物种: 24.7%), LDMC较低(个体: 17.0%, 物种: 14.1%); 当年生叶片MLA、LDMC和SLA的变异系数均高于往年生叶片; 2)往年生叶MLA显著大于当年生叶(t = -38.53, p < 0.001), 往年生叶SLA显著小于当年生叶(t = 45.30, p < 0.001), 往年生叶LDMC显著大于当年生叶(t = -9.71, p < 0.001); 3)在个体水平, 当年生叶片MLA、SLA和LDMC值分别解释了往年生叶片MLA、SLA和LDMC变异的86%、48%和41%; 在物种水平, 当年生叶片MLA、SLA和LDMC值分别解释了往年生叶片MLA、SLA和LDMC变异的97%、83%和85%; 4) SLA在异龄叶间的变化表明, 与往年生叶片相比, 投资相同干物质, 当年生叶片可形成较大的叶面积, 其叶片面积建成消耗较小。研究认为, 植物叶性状在异龄叶间具有较大的变异性和关联性, 叶面积形成过程中生物量建成与消耗的协调可能影响植物叶片的发育。     

Leaf traits indicate survival strategies among 42 dominant plant species in a dry, sandy habitat, China

The objective of this paper was to assess the congruency of leaf traits and soil characteristics and to analyze the survival strategies of different plant functional types in response to drought and nutrient-poor environ-ments in the southeastern Ke'erqin Sandy Lands in China. Six leaf traits-leaf thickness (TH), density (DN), specific leaf area (SLA), leaf dry weight to fresh weight ratio (DW/ FW), leaf N concentration (Nmass), and N resorption efficiency (NREmass)-of 42 plant species were investi-gated at four sites. The correlations between leaf traits and soil characteristics-organic C (OC), total N (TN), total P (TP), and soil moisture (SM)-were examined. We found that the six leaf traits across all the 42 species showed large variations and that DW/FW was negatively correlated with OC, TN, TP, and SM (P<0.05), while other leaf traits showed no significant correlations with soil characteristics. To find the dissimilarity to accommodate environment, a hierarchical agglomerative clustering analysis was made of all the species. All the species clustered into three groups except the Scutellaria baicalensis. Species of group Ⅲ might be most tolerant of an arid environment, and species of group Ⅱ might avoid nutrient stress in the nutrient-poor environment, while group Ⅰ was somewhat intermediate. Therefore, species from the different groups may be selected for use in vegetation restoration of different sites based on soil moisture and nutrient conditions.     

The profile distribution of total and extractable boron in cacao-growing soils in southwestern Nigeria

The profile distribution of total and extractable B was determined in 16 Nigerian cacao-growing soil profiles formed from different parent materials. Total B for all soils ranged from 8 to 54μgg⁻¹ with a mean of 24μgg⁻¹. The soils formed from sandstones in the rainforest zone contained higher amounts of total B than soils derived from basement complex. Boron extractable in hot water, in 0.1% CaCl₂, and in 1N NH₄OAc varied from 0.13 to 1.38, 0.44 to 1.20 0.03 to 0.56μgg⁻¹ respectively. The corresponding means were 0.66, 0.75 and 0.27μgg⁻¹ B. Soils on metamorphic rocks gave the highest values. All extractable B values were related to organic matter while only CaCl₂-extractable B correlated with total B. Generally total and extractable B values were higher in the top soils than in the subsoils.     

Spider web guilds in cacao agroforestry – comparing tree,plot and landscape‐scale management

Aim Owing to their role as insect predators, web‐building spiders can be important biological control agents within agricultural systems. In complex tropical agroecosystems such as agroforests, management determines plant architecture, vegetation composition and associated ant density, but little is known on how these attributes, together with landscape context, determine spider web density. We hypothesized that all three spatial scales and the presence of Philidris ants significantly contribute to the explanation of spider web density with web types being differently affected. Location In 42 differently managed cacao agroforestry systems in Sulawesi, Indonesia. Methods We surveyed the distribution of five spider‐web types on 420 cacao trees to determine how these relate to habitat variables and a numerically dominant ant species at three different spatial scales, comparing tree, plot and landscape features. We fitted linear mixed‐effects model, selected the best model subset using information‐theoretic criteria and calculated the model‐averaged estimates. We used non‐metric multidimensional scaling (NMDS) to determine and visualize guild level responses to the effects of the tree, plot and landscape‐scale variables. Results The five spider guilds preferred different features of cacao tree architecture. Most frequently recorded webs belonged to the line‐ and orb‐web type. At the tree scale, overall web density was positively related to canopy openness. At the plot scale, a higher number of shade trees was related to a higher web density. At the landscape scale, the altitude determined the distribution patterns of web‐building spiders. Presence of Philidris ants was positively associated with density of orb webs, while no pattern was found for other web types. Main conclusions Results suggest spider web density could be increased by pruning of cacao trees while keeping shade trees at high density in cacao plots. The results emphasize the need to consider scale dependency of crop management and web‐guild‐specific responses that may be related to different functional roles of spiders as a high‐density predator group in agroforestry.     

Micropropagation of <Emphasis Type="Italic">Theobroma cacao</Emphasis> L. using somatic embryo-derived plants

Summary A micropropagation protocol was developed using cacao somatic embryo-derived plant as a source for nodal and apical stem explants, and apical microcuttings. Microcuttings were efficiently rooted and developed into plantlets. Axillary meristems within the remaining decapitated plantlets subsequently developed and were used for production of additional microcuttings, with an average 2.4 growing shoots per decapitated stem. The remaining plantelts were maintained as microcutting stock plants. When nodal stem explants were cultured on thidiazuron medium, axillary buds proliferated and developed into shoots, which were excised and rooted. However, the efficiency of this method is lower than rooting of apical microcuttings harvested directly from stock plants. During root induction, short treatment with indole-3-butyric acid (IBA) increased the total percentage of rooted microcuttings up to 89%. Longer exposures to IBA increased the average number of roots per microcutting (from 1.7 to 5.2). Plant acclimatization after rooting was achieved with an average success of 87%. During several months of growth in the greenhouse, the micropropagated plants developed functional taproots. Currently, cocoa plants produced by this micropropagation method have been successfully acclimated to field conditions in Ivory Coast, Ghana, and Saint Lucia.     

Photosynthetic photon flux density,carbon dioxide concentration,and vapor pressure deficit effects on photosynthesis in cacao seedlings

Independent short-term effects of photosynthetic photon flux density (PPFD) of 50–400 μmol m⁻² s⁻¹, external CO₂ concentration (C _a) of 85–850 cm³ m⁻³, and vapor pressure deficit (VPD) of 0.9–2.2 kPa on net photosynthetic rate (P _N), stomatal conductance (g _s), leaf internal CO₂ concentration (C _i), and transpiration rates (E) were investigated in three cacao genotypes. In all these genotypes, increasing PPFD from 50 to 400 μmol m⁻² s⁻¹ increased P _N by about 50 %, but further increases in PPFD up to 1 500 μmol m⁻² s⁻¹ had no effect on P _N. Increasing C _a significantly increased P _N and C _i while g _s and E decreased more strongly than in most trees that have been studied. In all genotypes, increasing VPD reduced P _N, but the slight decrease in g _s and the slight increase in C _i with increasing VPD were non-significant. Increasing VPD significantly increased E and this may have caused the reduction in P _N. The unusually small response of g _s to VPD could limit the ability of cacao to grow where VPD is high. There were no significant differences in gas exchange characteristics (g _s, C _i, E) among the three cacao genotypes under any measurement conditions.