Phenology, growth and physiological adjustments of oil palm (Elaeis guineensis) to sink limitation induced by fruit pruning.

BACKGROUND AND AIMS: Despite its simple architecture and small phenotypic plasticity, oil palm has complex phenology and source-sink interactions. Phytomers appear in regular succession but their development takes years, involving long lag periods between environmental influences and their effects on sinks. Plant adjustments to resulting source-sink imbalances are poorly understood. This study investigated oil palm adjustments to imbalances caused by severe fruit pruning. METHODS: An experiment with two treatments (control and complete fruit pruning) during 22 months in 2006-2008) and six replications per treatment was conducted in Indonesia. Phenology, growth of above-ground vegetative and reproductive organs, leaf morphology, inflorescence sex differentiation, dynamics of non-structural carbohydrate reserves and light-saturated net photosynthesis (A(max)) were monitored. KEY RESULTS: Artificial sink limitation by complete fruit pruning accelerated development rate, resulting in higher phytomer, leaf and inflorescence numbers. Leaf size and morphology remained unchanged. Complete fruit pruning also suppressed the abortion of male inflorescences, estimated to be triggered at about 16 months before bunch maturity. The number of female inflorescences increased after an estimated lag of 24-26 months, corresponding to time from sex differentiation to bunch maturity. The most important adjustment process was increased assimilate storage in the stem, attaining nearly 50 % of dry weight in the stem top, mainly as starch, whereas glucose, which in controls was the most abundant non-structural carbohydrate stored in oil palm, decreased. CONCLUSIONS: The development rate of oil palm is in part controlled by source-sink relationships. Although increased rate of development and proportion of female inflorescences constituted observed adjustments to sink limitation, the low plasticity of plant architecture (constant leaf size, absence of branching) limited compensatory growth. Non-structural carbohydrate storage was thus the main adjustment process.

Phenology and growth adjustments of oil palm (Elaeis guineensis) to photoperiod and climate variability.

BACKGROUND AND AIMS: Oil palm flowering and fruit production show seasonal maxima whose causes are unknown. Drought periods confound these rhythms, making it difficult to analyse or predict dynamics of production. The present work aims to analyse phenological and growth responses of adult oil palms to seasonal and inter-annual climatic variability. METHODS: Two oil palm genotypes planted in a replicated design at two sites in Indonesia underwent monthly observations during 22 months in 2006-2008. Measurements included growth of vegetative and reproductive organs, morphology and phenology. Drought was estimated from climatic water balance (rainfall - potential evapotranspiration) and simulated fraction of transpirable soil water. Production history of the same plants for 2001-2005 was used for inter-annual analyses. KEY RESULTS: Drought was absent at the equatorial Kandista site (0 degrees 55'N) but the Batu Mulia site (3 degrees 12'S) had a dry season with variable severity. Vegetative growth and leaf appearance rate fluctuated with drought level. Yield of fruit, a function of the number of female inflorescences produced, was negatively correlated with photoperiod at Kandista. Dual annual maxima were observed supporting a recent theory of circadian control. The photoperiod-sensitive phases were estimated at 9 (or 9 + 12 x n) months before bunch maturity for a given phytomer. The main sensitive phase for drought effects was estimated at 29 months before bunch maturity, presumably associated with inflorescence sex determination. CONCLUSION: It is assumed that seasonal peaks of flowering in oil palm are controlled even near the equator by photoperiod response within a phytomer. These patterns are confounded with drought effects that affect flowering (yield) with long time-lag. Resulting dynamics are complex, but if the present results are confirmed it will be possible to predict them with models.

Role of transitory carbon reserves during adjustment to climate variability and source-sink imbalances in oil palm (Elaeis guineensis).

Oil palm (Elaeis guineensis Jacq.) is a perennial, tropical, monocotyledonous plant characterized by simple architecture and low phenotypic plasticity, but marked by long development cycles of individual phytomers (a pair of one leaf and one inflorescence at its axil). Environmental effects on vegetative or reproductive sinks occur with various time lags depending on the process affected, causing source-sink imbalances. This study investigated how the two instantaneous sources of carbon assimilates, CO(2) assimilation and mobilization of transitory non-structural carbohydrate (NSC) reserves, may buffer such imbalances. An experiment was conducted in Indonesia during a 22-month period (from July 2006 to May 2008) at two contrasting locations (Kandista and Batu Mulia) using two treatments (control and complete fruit pruning treatment) in Kandista. Measurements included leaf gas exchange, dynamics of NSC reserves and dynamics of structural aboveground vegetative growth (SVG) and reproductive growth. Drought was estimated from a simulated fraction of transpirable soil water. The main sources of variation in source-sink relationships were (i) short-term reductions in light-saturated leaf CO(2) assimilation rate (A(max)) during seasonal drought periods, particularly in Batu Mulia; (ii) rapid responses of SVG rate to drought; and (iii) marked lag periods between 16 and 29 months of environmental effects on the development of reproductive sinks. The resulting source-sink imbalances were buffered by fluctuations in NSC reserves in the stem, which mainly consisted of glucose and starch. Starch was the main buffer for sink variations, whereas glucose dynamics remained unexplained. Even under strong sink limitation, no negative feedback on A(max) was observed. In conclusion, the different lag periods for environmental effects on assimilate sources and sinks in oil palm are mainly buffered by NSC accumulation in the stem, which can attain 50% (dw:dw) in stem tops. The resulting dynamics of growth and production are complex because several dozen phytomers of different phenological ages develop at any given time and interact with a common pool of reserves.

Fruit development, not GPP, drives seasonal variation in NPP in a tropical palm plantation.

We monitored seasonal variations in net primary production (NPP), estimated by allometric equations from organ dimensions, gross primary production (GPP), estimated by the eddy covariance method, autotrophic respiration (R(a)), estimated by a model, and fruit production in a coconut (Cocos nucifera L.) plantation located in the sub-tropical South Pacific archipelago of Vanuatu. Net primary production of the vegetative compartments of the trees accumulated steadily throughout the year. Fruits accounted for 46% of tree NPP and showed large seasonal variations. On an annual basis, the sum of estimated NPP (16.1 Mg C ha(-1) year(-1)) and R(a) (24.0 Mg C ha(-1) year(-1)) for the ecosystem (coconut trees and herbaceous understory) closely matched GPP (39.0 Mg C ha(-1) year(-1)), suggesting adequate cross-validation of annual C budget methods. However, seasonal variations in NPP + R(a) were smaller than the seasonal variations in GPP, and maximum tree NPP occurred 6 months after the midsummer peak in GPP and solar radiation. We propose that this discrepancy reflects seasonal variation in the allocation of dry mass to carbon reserves and new plant tissue, thus affecting the allometric relationships used for estimating NPP.

Whole-plant adjustments in coconut (Cocos nucifera) in response to sink-source imbalance.

Coconut (Cocos nucifera L.) is a perennial tropical monocotyledon that produces fruit continuously. The physiological function of the large amounts of sucrose stored in coconut stems is unknown. To test the hypothesis that reserve storage and mobilization enable the crop to adjust to variable sink-source relationships at the scale of the whole plant, we investigated the dynamics of dry matter production, yield and yield components, and concentrations of nonstructural carbohydrate reserves in a coconut plantation on Vanuatu Island in the South Pacific. Two treatments were implemented continuously over 29 months (April 2002 to August 2004): 50% leaf pruning (to reduce the source) and 100% fruit and inflorescence pruning (to reduce the sink). The pruning treatments had little effect on carbohydrate reserves because they affected only petioles, not the main reserve pool in the stem. Both pruning treatments greatly reduced dry matter production of the reproductive compartment, but vegetative growth and development were negligibly affected by treatment and season. Leaf pruning increased radiation-use efficiency (RUE) initially, and fruit pruning greatly reduced RUE throughout the experiment. Changes in RUE were negatively correlated with leaflet soluble sugar concentration, indicating feedback inhibition of photosynthesis. We conclude that vegetative development and growth of coconut show little phenotypic plasticity, assimilate demand for growth being largely independent of a fluctuating assimilate supply. The resulting sink-source imbalances were partly compensated for by transitory reserves and, more importantly, by variable RUE in the short term, and by adjustment of fruit load in the long term. Possible physiological mechanisms are discussed, as well as modeling concepts that may be applied to coconut and similar tree crops.