Phenotypic and genetic dissection of component traits for early vigour in rice using plant growth modelling, sugar content analyses and association mapping.

Early vigour of rice, defined as seedling capacity to accumulate shoot dry weight (SDW) rapidly, is a complex trait. It depends on a genotype propensity to assimilate, store, and/or use non-structural carbohydrates (NSC) for producing large and/or numerous leaves, involving physiological trade-offs in the expression of component traits and, possibly, physiological and genetic linkages. This study explores a plant-model-assisted phenotyping approach to dissect the genetic architecture of rice early vigour, applying the Genome Wide Association Study (GWAS) to morphological and NSC measurements, as well as fitted parameters for the functional-structural plant model, Ecomeristem. Leaf size, number, SDW, and source-leaf NSC concentration were measured on a panel of 123 japonica accessions. The data were used to estimate Ecomeristem genotypic parameters driving organ appearance rate, size, and carbon dynamics. GWAS was performed based on 12 221 single-nucleotide polymorphisms (SNP). Twenty-three associations were detected at P <1×10(-4) and 64 at P <5×10(-4). Associations for NSC and model parameters revealed new regions related to early vigour that had greater significance than morphological traits, providing additional information on the genetic control of early vigour. Plant model parameters were used to characterize physiological and genetic trade-offs among component traits. Twelve associations were related to loci for cloned genes, with nine related to organogenesis, plant height, cell size or cell number. The potential use of these associations as markers for breeding is discussed.

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.

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.

Variability of phyllochron, plastochron and rate of increase in height in photoperiod-sensitive sorghum varieties.

BACKGROUND AND AIMS: West African sorghum (Sorghum bicolor) varieties are generally highly photoperiod-sensitive, which is a necessary adaptation to the variable onset date of the rainy season and the variable dates of sowing in the savannah zone. Depending on sowing date, plants can produce from 12 to >40 leaves on the main culm, with height varying from 1 m to more than 5 m. The present study aimed to better understand the complex phenology of these variables. METHODS: A 2-year series of monthly sowings of three West African sorghum varieties was conducted near Bamako, Mali. Drought stress was avoided by supplemental irrigation. Rate of initiation of primordia at the stem apex was recorded, together with rate of leaf emergence and increase in plant height. KEY RESULTS: Leaf initiation and appearance rates (plastochron(-1) and phyllochron(-1)) were constant for a given sowing date in cases where less than 20 leaves were produced (generally observed with late sowing dates). In contrast, rates were bilinear for early sowing dates, for which plants produced more than 20 leaves. The secondary rates, which occurred from the 20th leaf onwards, were only half of the initial rate. Plastochron and phyllochron showed large variations among sowing dates, and were correlated with the rate of plant height increase. The initial plastochron and phyllochron were positively correlated with soil temperature and negatively correlated with both day length and day-to-day change of day length prevailing at plant emergence, but these factors explained only half of the variation observed. CONCLUSIONS: Although they belong to different genetic groups and have different height and photoperiod sensitivity, the three varieties studied exhibited similar response patterns of development rates among phenological phases and seasons, with the local landrace showing the greatest variation due to its longer vegetative phase and longer stem internodes. The possible adaptive advantages in African savannah environments of bilinear development rates and the associated limitation in height increase are discussed.

Orchestration of transpiration, growth and carbohydrate dynamics in rice during a dry-down cycle.

The regulation of carbohydrate metabolism and source-sink relationships among organs play a key role in plant adaptation to drought. This study aimed at characterising the dynamics of transpiration, development, growth and carbon metabolism, as well as the expression of invertase genes, in response to drought during a dry-down cycle. Three 1-month experiments were conducted in controlled environment using the rice genotype IR64 (Oryza sativa L., indica). Plant leaf relative transpiration and expansion rates decreased linearly when fraction of transpirable soil water (FTSW) dropped below 0.66 and 0.58, respectively. Hexose and starch concentration responses to FTSW in a given organ were generally linear and opposite: in source leaves, hexose concentration increased and starch decreased, and vice versa in sink leaves and roots. Sucrose remained constant in source leaves and increased slightly in sink leaves. Starch reserves built up during stress in sink organs were rapidly mobilised upon rewatering, indicating its involvement in a mechanism to ensure recovery. Expression of cell-wall and vacuolar invertase genes under stress increased in sink leaves, interpreted as a mechanism to maintain sink activity (cell wall) and osmotic adjustment (vacuolar). It is concluded that carbohydrate metabolism in sink organs under drought is highly regulated, and important for stress adaptation.

Effect of abscisic acid on the transport of assimilates in barley.

The effect of abscisic acid (ABA) on assimilate transport in barley was investigated in two parallel experiments. First, the effect upon [(14)C]sucrose transport from the flag leaf to the ear of a single ABA application made at different stages of growth of the fruits was investigated; the effect was measured 24 h after treatment. Second, the effect of a single application of ABA made at the same stages of growth as above on grain weight of the mature plant was investigated. In both types of experiments ABA was applied once to the ear of different plants as an aqueous solution (10(-3)-10(-5) M), one to five weeks after anthesis. [(14)C] sucrose was applied by means of agar blocks. Parallel to these experiments, the endogenous content of ABA was investigated in the developing grains. When ears were treated with ABA two or four weeks after anthesis, an increase of up to 70% in the (14)C-transport from the flag leaf to the ear was observed within a 24-h period after treatment (short duration experiments). At these growth stages the endogenous concentrations of ABA were low. In sharp contrast, ABA, especially in a concentration of 10(-3) M, decreased (14)C-import from the flag leaf when applied three weeks after anthesis. At this stage the endogenous ABA content had reached its maximum. Long duration experiments with a single application of ABA to the car two weeks after anthesis resulted in a marked increase of weight per thousand kernels. ABA applications made earlier or later than two weeks after anthesis either reduced the grain weight or had no effect. It is concluded that ABA is involved in the regulation of assimilate transport from the leaves to the grains, possibly by influencing the unloading of sieve tubes in the ears. Promotion or inhibition of assimilate import by exogenously applied ABA may depend on the developmental stage of the grains and on the endogenous ABA level.