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1.  Genetic analysis of potassium use efficiency in Brassica oleracea 
Annals of Botany  2009;105(7):1199-1210.
Background and Aims
Potassium (K) fertilizers are used in intensive and extensive agricultural systems to maximize production. However, there are both financial and environmental costs to K-fertilization. It is therefore important to optimize the efficiency with which K-fertilizers are used. Cultivating crops that acquire and/or utilize K more effectively can reduce the use of K-fertilizers. The aim of the present study was to determine the genetic factors affecting K utilization efficiency (KUtE), defined as the reciprocal of shoot K concentration (1/[K]shoot), and K acquisition efficiency (KUpE), defined as shoot K content, in Brassica oleracea.
Genetic variation in [K]shoot was estimated using a structured diversity foundation set (DFS) of 376 accessions and in 74 commercial genotypes grown in glasshouse and field experiments that included phosphorus (P) supply as a treatment factor. Chromosomal quantitative trait loci (QTL) associated with [K]shoot and KUpE were identified using a genetic mapping population grown in the glasshouse and field. Putative QTL were tested using recurrent backcross substitution lines in the glasshouse.
Key Results
More than two-fold variation in [K]shoot was observed among DFS accessions grown in the glasshouse, a significant proportion of which could be attributed to genetic factors. Several QTL associated with [K]shoot were identified, which, despite a significant correlation in [K]shoot among genotypes grown in the glasshouse and field, differed between these two environments. A QTL associated with [K]shoot in glasshouse-grown plants (chromosome C7 at 62·2 cM) was confirmed using substitution lines. This QTL corresponds to a segment of arabidopsis chromosome 4 containing genes encoding the K+ transporters AtKUP9, AtAKT2, AtKAT2 and AtTPK3.
There is sufficient genetic variation in B. oleracea to breed for both KUtE and KUpE. However, as QTL associated with these traits differ between glasshouse and field environments, marker-assisted breeding programmes must consider carefully the conditions under which the crop will be grown.
PMCID: PMC2887060  PMID: 19815571
Arabidopsis; Brassica oleracea; genetics; potassium (K); potassium use efficiency (KUE); quantitative trait loci (QTL); shoot
2.  New insights into ferritin synthesis and function highlight a link between iron homeostasis and oxidative stress in plants 
Annals of Botany  2009;105(5):811-822.
Iron is an essential element for both plant productivity and nutritional quality. Improving plant iron content was attempted through genetic engineering of plants overexpressing ferritins. However, both the roles of these proteins in plant physiology, and the mechanisms involved in the regulation of their expression are largely unknown. Although the structure of ferritins is highly conserved between plants and animals, their cellular localization differs. Furthermore, regulation of ferritin gene expression in response to iron excess occurs at the transcriptional level in plants, in contrast to animals which regulate ferritin expression at the translational level.
In this review, an overview of our knowledge of bacterial and mammalian ferritin synthesis and functions is presented. Then the following will be reviewed: (a) the specific features of plant ferritins; (b) the regulation of their synthesis during development and in response to various environmental cues; and (c) their function in plant physiology, with special emphasis on the role that both bacterial and plant ferritins play during plant–bacteria interactions. Arabidopsis ferritins are encoded by a small nuclear gene family of four members which are differentially expressed. Recent results obtained by using this model plant enabled progress to be made in our understanding of the regulation of the synthesis and the in planta function of these various ferritins.
Studies on plant ferritin functions and regulation of their synthesis revealed strong links between these proteins and protection against oxidative stress. In contrast, their putative iron-storage function to furnish iron during various development processes is unlikely to be essential. Ferritins, by buffering iron, exert a fine tuning of the quantity of metal required for metabolic purposes, and help plants to cope with adverse situations, the deleterious effects of which would be amplified if no system had evolved to take care of free reactive iron.
PMCID: PMC2859905  PMID: 19482877
Iron; bacterioferritins; ferritins; oxidative stress; iron storage; seeds; pathogens; nutrition
3.  Nitric oxide and frataxin: two players contributing to maintain cellular iron homeostasis 
Annals of Botany  2009;105(5):801-810.
Nitric oxide (NO) is a signalling and physiologically active molecule in animals, plants and bacteria. The specificity of the molecular mechanism(s) involved in transducing the NO signal within and between cells and tissues is still poorly understood. NO has been shown to be an emerging and potent signal molecule in plant growth, development and stress physiology. The NO donor S-nitrosoglutathion (GSNO) was shown to be a biologically active compound in plants and a candidate for NO storage and/or mobilization between plant tissues and cells. NO has been implicated as a central component in maintaining iron bioavailavility in plants.
Scope and Conclusions
Iron is an essential nutrient for almost all organisms. This review presents an overview of the functions of NO in iron metabolism in animals and discusses how NO production constitutes a key response in plant iron sensing and availability. In plants, NO drives downstream responses to both iron deficiency and iron overload. NO-mediated improvement of iron nutrition in plants growing under iron-deficient conditions represents a powerful tool to cope with soils displaying low iron availability. An interconversion between different redox forms based on the iron and NO status of the plant cells might be the core of a metabolic process driving plant iron homeostasis. Frataxin, a recently identified protein in plants, plays an important role in mitochondria biogenesis and in maintaining mitochondrial iron homeostasis. Evidence regarding the interaction between frataxin, NO and iron from analysis of frataxin knock-down Arabidopsis thaliana mutants is reviewed and discussed.
PMCID: PMC2859906  PMID: 19556267
Nitric oxide; iron homeostasis; frataxin; strategy I; strategy II; dinitrosyl iron complexes; oxidative stress
4.  A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration 
Annals of Botany  2009;105(3):431-442.
Background and Aims
Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO2. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO2 and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.
An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO2 and H2O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO2 to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO2.
Key Results and Conclusion
In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO2, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO2; (b) the new formulation correctly predicts the condition under which CO2-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.
PMCID: PMC2826246  PMID: 19995810
Economics of gas exchange; free air CO2 enrichment; marginal water-use efficiency; photosynthesis; Pinus taeda; stomatal conductance; stomatal optimization
5.  Effects of phosphorus supply on growth, phosphate concentration and cluster-root formation in three Lupinus species 
Annals of Botany  2009;105(3):365-374.
Background and Aims
In some lupin species, phosphate deficiency induces cluster-root formation, which enhances P uptake by increasing root surface area and, more importantly, the release of root exudates which enhances P availability.
Three species of Lupinus, L. albus, L. atlanticus and L. micranthus, with inherently different relative growth rates were cultivated under hydroponics in a greenhouse at four phosphate concentrations (1, 10, 50 and 150 µm) to compare the role of internal P in regulating cluster-root formation.
Key Results
The highest growth rate was observed in L. atlanticus, followed by L. albus and L. micranthus. At 1 µm P, cluster-root formation was markedly induced in all three species. The highest P uptake and accumulation was observed in L. micranthus, followed by L. atlanticus and then L. albus. Inhibition of cluster-root formation was severe at 10 µm P in L. atlanticus, but occurred stepwise with increasing P concentration in the root medium in L. albus.
In L. atlanticus and L. albus cluster-root formation was suppressed by P treatments above 10 µm, indicating a P-inducible regulating system for cluster-root formation, as expected. By contrast, production of cluster roots in L. micranthus, in spite of a high internal P concentration, indicated a lower sensitivity to P status, which allowed P-toxicity symptoms to develop.
PMCID: PMC2826247  PMID: 20037142
Cluster roots; lupin; Lupinus; phosphate nutrition; toxicity; uptake
6.  Evolution and history of grapevine (Vitis vinifera) under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars 
Annals of Botany  2009;105(3):443-455.
Background and Aims
In spite of the abundance of archaeological, bio-archaeological, historical and genetic data, the origins, historical biogeography, identity of ancient grapevine cultivars and mechanisms of domestication are still largely unknown. Here, analysis of variation in seed morphology aims to provide accurate criteria for the discrimination between wild grapes and modern cultivars and to understand changes in functional traits in relation to the domestication process. This approach is also used to quantify the phenotypic diversity in the wild and cultivated compartments and to provide a starting point for comparing well-preserved archaeological material, in order to elucidate the history of grapevine varieties.
Geometrical analysis (elliptic Fourier transform method) was applied to grapevine seed outlines from modern wild individuals, cultivars and well-preserved archaeological material from southern France, dating back to the first to second centuries.
Key Results and Conclusions
Significant relationships between seed shape and taxonomic status, geographical origin (country or region) of accessions and parentage of varieties are highlighted, as previously noted based on genetic approaches. The combination of the analysis of modern reference material and well-preserved archaeological seeds provides original data about the history of ancient cultivated forms, some of them morphologically close to the current ‘Clairette’ and ‘Mondeuse blanche’ cultivars. Archaeobiological records seem to confirm the complexity of human contact, exchanges and migrations which spread grapevine cultivation in Europe and in Mediterranean areas, and argue in favour of the existence of local domestication in the Languedoc (southern France) region during Antiquity.
PMCID: PMC2826248  PMID: 20034966
Domestication syndrome; origin of cultivars; Vitis vinifera; seed; elliptic Fourier transforms
8.  Structure and function of plants 
Annals of Botany  2009;105(2):304-vii.
PMCID: PMC2814747
9.  Plant biology 
Annals of Botany  2009;105(2):679-viii.
PMCID: PMC2814748
10.  Architectural strategies of Cornus sericea, a native but invasive shrub of Southern Quebec, Canada, under an open or a closed canopy 
Annals of Botany  2009;105(2):205-220.
Background and Aims
Qualitative and quantitative studies of the pattern of invasive plant development is considered a key aspect in understanding invasiveness. An architectural analysis was therefore performed in order to understand the relationship between shoot architecture and invasiveness in red-osier dogwood, Cornus sericea (Cornaceae).
The structural and ontogenic characteristics of individuals in invading and non-invading populations in the native range of the species were compared to test the implication of developmental plasticity on invasiveness.
Key Results and Conclusions
The results show that the shrub has a modular architecture governed by strong developmental rules. Cornus sericea is made up of two levels of organization, each with its own intrinsic sequence of differentiation. These intrinsic mechanisms were used as a framework for comparison and it was found that, in response to the light environment, developmental plasticity was elevated, resulting in two architectural strategies. This developmental plasticity concerns the growth direction and the size of the modules, the speed of their time-course changes, their branching and flowering. Under an open canopy, C. sericea rapidly develops large vertical structures and abundant flowering. This strategy leads the plant to be invasive by excluding competitors and disseminating in the landscape. In the understorey, C. sericea slowly develops long horizontal structures which creep across the soil surface, while assimilating structures are poorly developed. This strategy does not lead to invasiveness but may allow the plant to survive in the understorey and reach sunny patches.
PMCID: PMC2814749  PMID: 19900945
Cornus sericea; Cornus stolonifera; red-osier dogwood; shrub; invasive; architecture; phenotypic plasticity; development; light; human disturbance
11.  Seasonal influences on carbohydrate metabolism in the CAM bromeliad Aechmea ‘Maya’: consequences for carbohydrate partitioning and growth 
Annals of Botany  2009;105(2):301-309.
Background and Aims
Photosynthetic plasticity in response to a range of environmental factors that include [CO2], water availability, light intensity and temperature, is ubiquitous among plants with crassulacean acid metabolism (CAM). The present study examined how seasonal changes in light availability, as experienced by greenhouse CAM crops in northern latitude regions, influence diel carboxylation patterns and impact on carbon gain and seasonal accumulation of biomass.
In the CAM bromeliad Aechmea ‘Maya’ integrated measurements of leaf gas exchange, diel metabolite dynamics (e.g. malate, soluble sugars and starch) and biomass accumulation were made four times a year, i.e. in winter, spring, summer and autumn.
Key Results
During the brighter seasons (spring and summer) daytime Phases II and IV were dominated by C4 carboxylation, whilst the higher diurnal uptake in the autumn and winter was characterized by equal contributions of both Rubisco and PEPC. As a consequence, net CO2 uptake showed a significant depression at the end of the day in the darker months when supplementary illumination was turned off. Remarkable seasonal consistency was found in the amount of storage reserves available for nocturnal carboxylation, a consequence of predominantly daytime export of carbohydrate in spring and summer whilst nocturnal export was the major sink for carbohydrate in autumn and winter.
Throughout the different seasons Aechmea ‘Maya’ showed considerable plasticity in the timing and magnitude of C3 and C4 carboxylation processes over the diel cycle. Under low PPFD (i.e. winter and autumn) it appears that there was a constraint on the amount of carbohydrate exported during the day in order to maintain a consistent pool of transient carbohydrate reserves. This gave remarkable seasonal consistency in the amount of storage reserves available at night, thereby optimizing biomass gain throughout the year. The data have important practical consequences for horticultural productivity of CAM plants and suggest a scenario for reconciling carbohydrate partitioning between competing sinks of nocturnal acidification and export for growth.
PMCID: PMC2814750  PMID: 19914918
Aechmea ‘Maya’; seasonal; CAM; bromeliad; carbon budgets; gas exchange; metabolite dynamics; PEPC; photoperiod extension; PPFD; photosynthetic plasticity; Rubisco
12.  Are the common assimilate pool and trophic relationships appropriate for dealing with the observed plasticity of grapevine development? 
Annals of Botany  2009;105(2):233-247.
Background and Aims
Models based on the consideration of plant development as the result of source–sink relationships between organs suffer from an inherent lack of quantification of the effect of trophic competition on organ growth processes. The ‘common assimilate pool theory’ underlying many such models is highly debatable.
Six experiments were carried out in a greenhouse and outdoors with two grapevine cultivars and with 12 management systems, resulting in different types of plant architecture. Ten variables were used to quantify the impact of variations in assimilate supply and topological distances between sources and sinks on organogenesis, morphogenesis and biomass growth.
Key Results
A hierarchy of the responses of these processes to variations in assimilate supply was identified. Organ size seemed to be independent of assimilate supply, whereas both organogenesis and biomass growth were affected by variations in assimilate supply. Lower levels of organ biomass growth in response to the depletion of assimilate supplies seemed to be the principal mechanism underlying the plasticity of plant development in different environments. Defoliation or axis ablation resulted in changes in the relationship between growth processes and assimilate supply, highlighting the influence of non-trophic determinants. The findings cast doubt on the relevance of ‘the common assimilate pool theory’ for modelling the development of grapevine.
The results of this study suggest new formalisms for increasing the ability of models to take plant plasticity into account. The combination of an ecophysiological model for morphogenesis taking environmental signals into account and a biomass driven model for organogenesis and biomass allocation taking the topological distances between the sources and the sinks into account appears to be a promising approach. Moreover, in order to simulate the impact of agronomic practices, it will be necessary to take into account the non-trophic determinants of plant development such as hormonal signaletics.
PMCID: PMC2814752  PMID: 19946042
Biomass growth; branching system; common assimilate pool; morphogenesis; organogenesis; source–sink; grapevine; Vitis vinifera
13.  Morphological evolution in the variable resin-producing Detarieae (Fabaceae): do morphological characters retain a phylogenetic signal? 
Annals of Botany  2009;105(2):311-325.
Background and Aims
Previous molecular phylogenetic studies disagree with the informal generic-level taxonomic groups based on morphology. In this study morphological characters in the caesalpinioid clade Detarieae are evaluated within a phylogenetic framework as a means of better understanding phylogenetic relationships and morphological evolution.
Morphological characters were observed and scored for representative species of Detarieae focusing on the resin-producing genera. Phylogenetic analyses were carried out with morphological characters alone and then combined with DNA sequences.
Key Results
Despite a high level of homoplasy, morphological data support several clades corresponding to those recovered in molecular phylogenetic analyses. The more strongly supported clades are each defined by at least one morphological synapomorphy. Several characters (e.g. apetaly) previously used to define informal generic groups evolved several times independently, leading to the differences observed with the molecular phylogenetic analyses. Although floral evolution is complex in Detarieae some patterns are recovered.
New informal taxonomic groupings are proposed based on the present findings. Floral evolution in the diverse Detarieae clade is characterized by a repeated tendency toward zygomorphy through the reduction of lateral petals and toward complete loss of petals.
PMCID: PMC2814753  PMID: 19939978
Caesalpinioideae; Detarieae; floral evolution; Leguminosae; morphology; phylogeny; resins; taxonomy
14.  Relationships between xylem vessel characteristics, calculated axial hydraulic conductance and size-controlling capacity of peach rootstocks 
Annals of Botany  2009;105(2):327-331.
Background and Aims
Previous studies indicate that the size-controlling capacity of peach rootstocks is associated with reductions of scion water potential during mid-day that are caused by the reduced hydraulic conductance of the rootstock. Thus, shoot growth appears to be reduced by decreases in stem water potential. The aim of this study was to investigate the mechanism of reduced hydraulic conductance in size-controlling peach rootstocks.
Anatomical measurements (diameter and frequency) of xylem vessels were determined in shoots, trunks and roots of three contrasting peach rootstocks grown as trees, each with different size-controlling characteristics: ‘Nemaguard’ (vigorous), ‘P30-135’ (intermediate vigour) and ‘K146-43’ (substantially dwarfing). Based on anatomical measurements, the theoretical axial xylem conductance of each tissue type and rootstock genotype was calculated via the Poiseuille–Hagen law.
Key Results
Larger vessel dimensions were found in the vigorous rootstock (‘Nemaguard’) than in the most dwarfing one (‘K146-43’) whereas vessels of ‘P30-135’ had intermediate dimensions. The density of vessels per xylem area in ‘Nemaguard’ was also less than in ‘P30-135’and ‘K146-43’. These characteristics resulted in different estimated hydraulic conductance among rootstocks: ‘Nemaguard’ had higher theoretical values followed by ‘P30-135’ and ‘K146-43’.
These data indicate that phenotypic differences in xylem anatomical characteristics of rootstock genotypes appear to influence hydraulic conductance capacity directly, and therefore may be the main determinant of dwarfing in these peach rootstocks.
PMCID: PMC2814754  PMID: 19939979
Prunus; rootstock; vessel diameter; hydraulic conductance; dwarfing; xylem anatomy; Poiseuille–Hagen
15.  Cytotype diversity and genome size variation in eastern Asian polyploid Cardamine (Brassicaceae) species 
Annals of Botany  2009;105(2):249-264.
Background and Aims
Intraspecific ploidy-level variation is an important aspect of a species' genetic make-up, which may lend insight into its evolutionary history and future potential. The present study explores this phenomenon in a group of eastern Asian Cardamine species.
Plant material was sampled from 59 localities in Japan and Korea, which were used in karyological (chromosome counting) and flow cytometric analyses. The absolute nuclear DNA content (in pg) was measured using propidium iodide and the relative nuclear DNA content (in arbitrary units) was measured using 4,6-diamidino-2-phenylindole fluorochrome.
Key Results
Substantial cytotype diversity was found, with strikingly different distribution patterns between the species. Two cytotypes were found in C. torrentis sensu lato (4x and 8x, in C. valida and C. torrentis sensu stricto, respectively), which displays a north–south geographical pattern in Japan. Hypotheses regarding their origin and colonization history in the Japanese archipelago are discussed. In Korean C. amaraeiformis, only tetraploids were found, and these populations may in fact belong to C. valida. C. yezoensis was found to harbour as many as six cytotypes in Japan, ranging from hexa- to dodecaploids. Ploidy levels do not show any obvious geographical pattern; populations with mixed ploidy levels, containing two to four cytotypes, are frequently observed throughout the range. C. schinziana, an endemic of Hokkaido, has hexa- and octoploid populations. Previous chromosome records are also revised, showing that they are largely based on misidentified material or misinterpreted names.
Sampling of multiple populations and utilization of the efficient flow cytometric approach allowed the detection of large-scale variation in ploidy levels and genome size variation attributable to aneuploidy. These data will be essential in further phylogenetic and evolutionary studies.
PMCID: PMC2814755  PMID: 20007978
Cardamine amaraeiformis; Cardamine schinziana; Cardamine torrentis; Cardamine valida; Cardamine yezoensis; cytogeography; DNA ploidy level; flow cytometry; Japan; Korea; polyploidy
16.  Stereological estimation of cell wall density of DR12 tomato mutant using three-dimensional confocal imaging 
Annals of Botany  2009;105(2):265-276.
Background and Aims
The cellular structure of fleshy fruits is of interest to study fruit shape, size, mechanical behaviour or sensory texture. The cellular structure is usually not observed in the whole fruit but, instead, in a sample of limited size and volume. It is therefore difficult to extend measurements to the whole fruit and/or to a specific genotype, or to describe the cellular structure heterogeneity within the fruit.
An integrated method is presented to describe the cellular structure of the whole fruit from partial three-dimensional (3D) observations, involving the following steps: (1) fruit sampling, (2) 3D image acquisition and processing and (3) measurement and estimation of relevant 3D morphological parameters. This method was applied to characterize DR12 mutant and wild-type tomatoes (Solanum lycopersicum).
Key Results
The cellular structure was described using the total volume of the pericarp, the surface area of the cell walls and the ratio of cell-wall surface area to pericarp volume, referred to as the cell-wall surface density. The heterogeneity of cellular structure within the fruit was investigated by estimating variations in the cell-wall surface density with distance to the epidermis.
The DR12 mutant presents a greater pericarp volume and an increase of cell-wall surface density under the epidermis.
PMCID: PMC2814756  PMID: 19952012
Stereology; 3D morphology estimation; tomato fruit; cellular structure; image analysis; confocal microscopy
17.  Interspecific correlates of plasticity in relative growth rate following a decrease in nitrogen availability 
Annals of Botany  2009;105(2):333-339.
Background and Aims
Nitrogen availability varies greatly over short time scales. This requires that a well-adapted plant modify its phenotype by an appropriate amount and at a certain speed in order to maximize growth and fitness. To determine how plastic ontogenetic changes in each trait interact and whether or not these changes are likely to maximize growth, ontogenetic changes in relative growth rate (RGR), net assimilation rate (NAR), specific leaf area (SLA) and root weight ratio (RWR), before and after a decrease in nitrogen supply, were studied in 14 herbaceous species.
Forty-four plants of each species were grown in hydroponic culture under controlled conditions in a control treatment where the supply of nitrogen remained constant at 1 mm, and in a stress treatment where the nitrogen supply was abruptly decreased from 1 to 0·01 mm during the growth period.
Key Results and Conclusions
In the treatment series, and in comparison with the control, NAR and RGR decreased, RWR increased, and SLA did not change except for the timing of ontogenetic change. Species having greater increases in the maximum rate of change in RWR also had smaller reductions in RGR; plasticity in RWR is therefore adaptive. In contrast, species which showed a greater decrease in NAR showed stronger reductions in RGR; plasticity in NAR is therefore not adaptive. Plasticity in RGR was not related to plasticity in SLA. There were no significant relationships among the plasticities in NAR, RWR or SLA. Potentially fast-growing species experienced larger reductions in RGR following the nitrogen reduction. These results suggest that competitive responses to interspecific competition for nitrogen might be positively correlated with the plasticity in the maximum rate of change in RWR in response to a reduction in nitrogen supply.
PMCID: PMC2814757  PMID: 19942581
Root weight ratio; net assimilation rate; nutrient stress; ontogenetic plasticity; specific leaf area
18.  The interface between metabolic and stress signalling 
Annals of Botany  2009;105(2):197-203.
It is becoming increasingly clear that stress and metabolic signalling networks interact and that this interaction is important in plant responses to herbivory, pathogen attack, drought, cold, heat and osmotic stresses including salinity. At the interface between these two major signalling systems are the hormone abscisic acid (ABA) and signalling factors including protein kinases and transcription factors.
This briefing reviews links between ABA, stress and sugar signalling, focusing on the roles of sucrose non-fermenting-1-related protein kinases (SnRKs), SnRK1-activating protein kinases (SnAKs), calcium-dependent protein kinases (CDPKs) and ABA response element binding proteins (AREBPs, which are transcription factors). Links between stress and nitrogen / amino acid signalling are also described, including the roles of a protein kinase called general control non-derepressible (GCN)-2 in regulating protein synthesis through phosphorylation of the α-subunit of translation initiation factor-2 (eIF2α) in response not only to decreases in amino acid levels but also to a range of stresses. Evidence of a link between sugar and amino acid signalling is explored, with nitrate reductase being a target for regulation by both SnRK1 and GCN2 through different mechanisms; possible links between SnRK1 and GCN2 via a pathway including the protein kinase target of rapamycin (TOR)-1 are described. The significance of these interactions to the concept of signalling networks as opposed to simple cascades and pathways, and the importance of the subject in the context of the predicted increase in severity and range of stresses that plants will have to withstand as a result of global climate change are discussed.
PMCID: PMC2814758  PMID: 20007158
ABA (abscisic acid); AREBP (ABA response element binding protein); CDPK (calcium-dependent protein kinase); climate change; GCN2 (general control non-derepressible 2); metabolic signalling; protein kinase; signalling networks; SnAK (SnRK1-activating protein kinase); SnRK (sucrose non-fermenting-1-related protein kinase); stress signalling; transcription factor
19.  Mechanosensing of stem bending and its interspecific variability in five neotropical rainforest species 
Annals of Botany  2009;105(2):341-347.
Background and Aims
In rain forests, sapling survival is highly dependent on the regulation of trunk slenderness (height/diameter ratio): shade-intolerant species have to grow in height as fast as possible to reach the canopy but also have to withstand mechanical loadings (wind and their own weight) to avoid buckling. Recent studies suggest that mechanosensing is essential to control tree dimensions and stability-related morphogenesis. Differences in species slenderness have been observed among rainforest trees; the present study thus investigates whether species with different slenderness and growth habits exhibit differences in mechanosensitivity.
Recent studies have led to a model of mechanosensing (sum-of-strains model) that predicts a quantitative relationship between the applied sum of longitudinal strains and the plant's responses in the case of a single bending. Saplings of five different neotropical species (Eperua falcata, E. grandiflora, Tachigali melinonii, Symphonia globulifera and Bauhinia guianensis) were subjected to a regimen of controlled mechanical loading phases (bending) alternating with still phases over a period of 2 months. Mechanical loading was controlled in terms of strains and the five species were subjected to the same range of sum of strains. The application of the sum-of-strain model led to a dose–response curve for each species. Dose–response curves were then compared between tested species.
Key Results
The model of mechanosensing (sum-of-strain model) applied in the case of multiple bending as long as the bending frequency was low. A comparison of dose–response curves for each species demonstrated differences in the stimulus threshold, suggesting two groups of responses among the species. Interestingly, the liana species B. guianensis exhibited a higher threshold than other Leguminosae species tested.
This study provides a conceptual framework to study variability in plant mechanosensing and demonstrated interspecific variability in mechanosensing.
PMCID: PMC2814759  PMID: 19995809
Mechanosensing; interspecific variability; trees; lianas; rain forest; neotropical species; bending; biomechanics; Bauhinia; Eperua; Symphonia; Tachigali
20.  The AUX1 LAX family of auxin influx carriers is required for the establishment of embryonic root cell organization in Arabidopsis thaliana 
Annals of Botany  2009;105(2):277-289.
Background and Aims
The root meristem of the Arabidopsis thaliana mature embryo is a highly organized structure in which individual cell shape and size must be regulated in co-ordination with the surrounding cells. The objective of this study was to determine the role of the AUX1 LAX family of auxin import carriers during the establishment of the embryonic root cell pattern.
The radicle apex of single and multiple aux1 lax mutant mature embryos was used to evaluate the effect of this gene family upon embryonic root organization and root cap size, cell number and cell size.
Key Results
It was demonstrated here that mutations within the AUX1 LAX family are associated with changes in cell pattern establishment in the embryonic quiescent centre and columella. aux1 lax mutants have a larger radicle root cap than the wild type and this is associated with a significant increase in the root-cap cell number, average cell size, or both. Extreme disorganization of the radicle apex was observed among quadruple aux1 lax1 lax2 lax3 mutant embryos, but not in single aux1 null or in lax1, lax2 and lax3 single mutants, indicating redundancy within the AUX1 LAX family.
It was determined that the AUX1 LAX family of auxin influx facilitators participates in the establishment of cell pattern within the apex of the embryonic root in a gene-redundant fashion. It was demonstrated that aux1 lax mutants are affected in cell proliferation and cell growth within the radicle tip. Thus AUX1 LAX auxin importers emerge as new players in morphogenetic processes involved in patterning during embryonic root formation.
PMCID: PMC2814760  PMID: 19952011
AUX1 LAX genes; auxin; Arabidopsis thaliana; embryogenesis; meristem; radicle development; cell pattern establishment
21.  Size matters for violent discharge height and settling speed of Sphagnum spores: important attributes for dispersal potential 
Annals of Botany  2009;105(2):291-300.
Background and Aims
Initial release height and settling speed of diaspores are biologically controlled components which are key to modelling wind dispersal. Most Sphagnum (peat moss) species have explosive spore liberation. In this study, how capsule and spore sizes affect the height to which spores are propelled were measured, and how spore size and spore number of discharged particles relate to settling speed in the aspherical Sphagnum spores.
Spore discharge and spore cloud development were filmed in a closed chamber (nine species). Measurements were taken from snapshots at three stages of cloud development. Settling speed of spores (14 species) and clusters were timed in a glass tube.
Key Results
The maximum discharge speed measured was 3·6 m s−1. Spores reached a maximum height of 20 cm (average: 15 cm) above the capsule. The cloud dimensions at all stages were related positively to capsule size (R2 = 0·58–0·65). Thus species with large shoots (because they have large capsules) have a dispersal advantage. Half of the spores were released as singles and the rest as clusters (usually two to four spores). Single spores settled at 0·84–1·86 cm s−1, about 52 % slower than expected for spherical spores with the same diameters. Settling speed displayed a positive curvilinear relationship with spore size, close to predictions by Stokes' law for spherical spores with 68 % of the actual diameters. Light-coloured spores settled slower than dark spores. Settling speed of spore clusters agrees with earlier studies. Effective spore discharge and small, slowly settling spores appear particularly important for species in forested habitats.
The spore discharge heights in Sphagnum are among the greatest for small, wind-dispersed propagules. The discharge heights and the slow settling of spores affect dispersal distances positively and may help to explain the wide distribution of most boreal Sphagnum species.
PMCID: PMC2814761  PMID: 20123930
Bryophyte; explosive capsules; long-distance dispersal; peatland; peat moss; settling speed; Sphagnum; spore; spore capsule; Sweden; terminal velocity; wind dispersal
22.  Pollen terminology. An illustrated handbook 
Annals of Botany  2009;105(2):264-ix.
PMCID: PMC2814762
24.  Convective gas flow development and the maximum depths achieved by helophyte vegetation in lakes 
Annals of Botany  2009;105(1):165-174.
Background and Aims
Convective gas flow in helophytes (emergent aquatic plants) is thought to be an important adaptation for the ability to colonize deep water. In this study, the maximum depths achieved by seven helophytes were compared in 17 lakes differing in nutrient enrichment, light attenuation, shoreline exposure and sediment characteristics to establish the importance of convective flow for their ability to form the deepest helophyte vegetation in different environments.
Convective gas flow development was compared amongst the seven species, and species were allocated to ‘flow absent’, ‘low flow’ and ‘high flow’ categories. Regression tree analysis and quantile regression analysis were used to determine the roles of flow category, lake water quality, light attenuation and shoreline exposure on maximum helophyte depths.
Key Results
Two ‘flow absent’ species were restricted to very shallow water in all lakes and their depths were not affected by any environmental parameters. Three ‘low flow’ and two ‘high flow’ species had wide depth ranges, but ‘high flow’ species formed the deepest vegetation far more frequently than ‘low flow’ species. The ‘low flow’ species formed the deepest vegetation most commonly in oligotrophic lakes where oxygen demands in sediments were low, especially on exposed shorelines. The ‘high flow’ species were almost always those forming the deepest vegetation in eutrophic lakes, with Eleocharis sphacelata predominant when light attenuation was low, and Typha orientalis when light attenuation was high. Depths achieved by all five species with convective flow were limited by shoreline exposure, but T. orientalis was the least exposure-sensitive species.
Development of convective flow appears to be essential for dominance of helophyte species in >0·5 m depth, especially under eutrophic conditions. Exposure, sediment characteristics and light attenuation frequently constrain them to a shallower depth than their flow capacity permits.
PMCID: PMC2794052  PMID: 19491087
Aeration; convective flow; exposure; helophytes; lakes; lakeshore vegetation; light attenuation; redox; regression tree; sediment motion; trophic state; waves
25.  Agricultural use of wetlands: opportunities and limitations 
Annals of Botany  2009;105(1):155-163.
Wetlands are species-rich habitats performing valuable ecosystem services such as flood protection, water quality enhancement, food chain support and carbon sequestration. Worldwide, wetlands have been drained to convert them into agricultural land or industrial and urban areas. A realistic estimate is that 50 % of the world's wetlands have been lost.
This paper reviews the relationship between wetlands and agriculture with the aim to identify the successes and failures of agricultural use in different types of wetlands, with reference to short-term and long-term benefits and issues of sustainability. It also addresses a number of recent developments which will lead to pressure to reclaim and destroy natural wetlands, i.e. the continuous need for higher production to feed an increasing world population and the increasing cultivation of energy crops. Finally, attention is paid to the development of more flood-tolerant crop cultivars.
Agriculture has been carried out in several types of (former) wetlands for millennia, with crop fields on river floodplain soils and rice fields as major examples. However, intensive agricultural use of drained/reclaimed peatlands has been shown to lead to major problems because of the oxidation and subsidence of the peat soil. This does not only lead to severe carbon dioxide emissions, but also results in low-lying land which needs to be protected against flooding. Developments in South-East Asia, where vast areas of tropical peatlands are being converted into oil palm plantations, are of great concern in this respect. Although more flood-tolerant cultivars of commercial crop species are being developed, these are certainly not suitable for cultivation in wetlands with prolonged flooding periods, but rather will survive relatively short periods of waterlogging in normally improved agricultural soils. From a sustainability perspective, reclamation of peatlands for agriculture should be strongly discouraged. The opportunities for agriculture in naturally functioning floodplains should be further investigated. The development and use of crop cultivars with an even stronger flood tolerance could form part of the sustainable use of such floodplain systems. Extensive use of wetlands without drastic reclamation measures and without fertilizer and pesticides might result in combinations of food production with other wetland services, with biodiversity remaining more or less intact. There is a need for research by agronomists and environmental scientists to optimize such solutions.
PMCID: PMC2794053  PMID: 19700447
Wetlands; sustainable agriculture; peat subsidence; floodplains; rice fields; water use; irrigation

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