Contrasting acclimation abilities of two dominant boreal conifers to elevated CO2 and temperature.

High latitude forests will experience large changes in temperature and CO2 concentrations this century. We evaluated the effects of future climate conditions on 2 dominant boreal tree species, Pinus sylvestris L. and Picea abies (L.) H. Karst, exposing seedlings to 3 seasons of ambient (430 ppm) or elevated CO2 (750 ppm) and ambient temperatures, a + 4 °C warming or a + 8 °C warming. Pinus sylvestris responded positively to warming: seedlings developed a larger canopy, maintained high net CO2 assimilation rates (Anet ), and acclimated dark respiration (Rdark ). In contrast, carbon fluxes in Picea abies were negatively impacted by warming: maximum rates of Anet decreased, electron transport was redirected to alternative electron acceptors, and thermal acclimation of Rdark was weak. Elevated CO2 tended to exacerbate these effects in warm-grown Picea abies, and by the end of the experiment Picea abies from the +8 °C, high CO2 treatment produced fewer buds than they had 3 years earlier. Treatments had little effect on leaf and wood anatomy. Our results highlight that species within the same plant functional type may show opposite responses to warming and imply that Picea abies may be particularly vulnerable to warming due to low plasticity in photosynthetic and respiratory metabolism.

Evaluating the involvement and interaction of abscisic acid and miRNA156 in the induction of anthocyanin biosynthesis in drought-stressed plants.

MAIN CONCLUSION: ABA is involved in anthocyanin synthesis through the regulation of microRNA156, augmenting the level of expression of anthocyanin synthesis-related genes and, therefore, increasing anthocyanin level. Drought stress is the main cause of agricultural crop loss in the world. However, plants have developed mechanisms that allow them to tolerate drought stress conditions. At cellular level, drought stress induces changes in metabolite accumulation, including increases in anthocyanin levels due to upregulation of the anthocyanin biosynthetic pathway. Recent studies suggest that the higher anthocyanin content observed under drought stress conditions could be a consequence of a rise in the abscisic acid (ABA) concentration. This plant hormone crosses the plasma membrane by specific transporters, and it is recognized at the cytosolic level by receptors known as pyrabactin resistance (PYR)/regulatory component of ABA receptors (PYR/RCARs) that regulate downstream components. In this review, we discuss the hypothesis regarding the involvement of ABA in the regulation of microRNA156 (miRNA156), which is upregulated as part of dehydration stress responsiveness in different species. The miRNA156 upregulation produces a greater level of anthocyanin gene expression, forming the multienzyme complex that will synthesize an increased level of anthocyanins at the cytosolic face of the rough endoplasmic reticulum (RER). After synthesis, anthocyanins are transported from the RER to the vacuole by two possible models of transport: (1) membrane vesicle-mediated transport, or (2) membrane transporter-mediated transport. Thus, the aim was to analyze the recent findings on synthesis, transport and the possible mechanism by which ABA could increase anthocyanin synthesis under drought stress conditions potentially throughout microRNA156 (miRNA156).

Photosynthetic acclimation, vernalization, crop productivity and 'the grand design of photosynthesis'.

Daniel Arnon first proposed the notion of a 'grand design of photosynthesis' in 1982 to illustrate the central role of photosynthesis as the primary energy transformer for all life on Earth. However, we suggest that this concept can be extended to the broad impact of photosynthesis not only in global energy transformation but also in the regulation of plant growth, development, survival and crop productivity through chloroplast redox signalling. We compare and contrast the role of chloroplast redox imbalance, measured as excitation pressure, in governing acclimation to abiotic stress and phenotypic plasticity. Although all photoautrophs sense excessive excitation energy through changes in excitation pressure, the response to this chloroplast redox signal is species dependent. Due to a limited capacity to adjust metabolic sinks, cyanobacteria and green algae induce photoprotective mechanisms which dissipate excess excitation energy at a cost of decreased photosynthetic performance. In contrast, terrestrial, cold tolerant plants such as wheat enhance metabolic sink capacity which leads to enhanced photosynthetic performance and biomass accumulation with minimal dependence on photoprotection. We suggest that the family of nuclear C-repeat binding transcription factors (CBFs) associated with the frost resistance locus, FR2, contiguous with the vernalization locus,VRN1, and mapped to chromosome 5A of wheat, may be critical components that link leaf chloroplast redox regulation to enhanced photosynthetic performance, the accumulation of growth-active gibberellins and the dwarf phenotype during cold acclimation prior to the vegetative to reproductive transition controlled by vernalization in winter cereals. Further genetic, molecular and biochemical research to confirm these links and to elucidate the molecular mechanism by which chloroplast redox modulation of CBF expression leads to enhanced photosynthetic performance is required. Because of the superior abiotic stress tolerance of cold tolerant winter wheat and seed yields that historically exceed those of spring wheat by 30-40%, we discuss the potential to exploit winter cereals for the maintenance or perhaps even the enhancement of cereal productivity under future climate change scenarios that will be required to feed a growing human population.

Evaluating the use of plant hormones and biostimulators in forage pastures to enhance shoot dry biomass production by perennial ryegrass (Lolium perenne L.).

Fertilisation of established perennial ryegrass forage pastures with nitrogen (N)-based fertilisers is currently the most common practice used on farms to increase pasture forage biomass yield. However, over-fertilisation can lead to undesired environmental impacts, including nitrate leaching into waterways and increased gaseous emissions of ammonia and nitrous oxide to the atmosphere. Additionally, there is growing interest from pastoral farmers to adopt methods for increasing pasture dry matter yield which use 'natural', environmentally safe plant growth stimulators, together with N-based fertilisers. Such plant growth stimulators include plant hormones and plant growth promotive microorganisms such as bacteria and fungi ('biostimulators', which may produce plant growth-inducing hormones), as well as extracts of seaweed (marine algae). This review presents examples and discusses current uses of plant hormones and biostimulators, applied alone or together with N-based fertilisers, to enhance shoot dry matter yield of forage pasture species, with an emphasis on perennial ryegrass.

Overexpression of gibberellin 20-oxidase1 from Pinus densiflora results in enhanced wood formation with gelatinous fiber development in a transgenic hybrid poplar.

Gibberellins (GAs) are important regulators of plant shoot biomass growth, and GA 20-oxidase (GA20ox) is one of the major regulatory enzymes in the GA biosynthetic pathway. Previously, we showed that the expression levels of a putative GA20ox1 (i.e., PdGA20ox1) in stem tissue of 3-month-old seedlings of 12 families of Pinus densiflora were positively correlated with stem diameter growth across those same families growing in an even-aged 32-year-old pine forest (Park EJ, Lee WY, Kurepin LV, Zhang R, Janzen L, Pharis RP (2015) Plant hormone-assisted early family selection in Pinus densiflora via a retrospective approach. Tree Physiol 35:86-94). To further investigate the molecular function of this gene in the stem wood growth of forest trees, we produced transgenic poplar lines expressing PdGA20ox1 under the control of the 35S promoter (designated as 35S::PdGA20ox1). By age 3 months, most of the 35S::PdGA20ox1 poplar trees were showing an exceptional enhancement of stem wood growth, i.e., up to fourfold increases in stem dry weight, compared with the nontransformed control poplar plants. Significant increases in endogenous GA1, its immediate precursor (GA20) and its catabolite (GA8) in elongating internode tissue accompanied the increased stem growth in the transgenic lines. Additionally, the development of gelatinous fibers occurred in vertically grown stems of the 35S::PdGA20ox1 poplars. An analysis of the cell wall monosaccharide composition of the 35S::PdGA20ox1 poplars showed significant increases in xylose and glucose contents, indicating a qualitative increase in secondary wall depositions. Microarray analyses led us to find a total of 276 probe sets that were upregulated (using threefold as a threshold) in the stem tissues of 35S::PdGA20ox1 poplars relative to the controls. 'Cell organization or biogenesis'- and 'cell wall'-related genes were overrepresented, including many of genes that are involved in cell wall modification. Several transcriptional regulators, which positively regulate cell elongation through GA signaling, were also upregulated. In contrast, genes involved in defense signaling were appreciably downregulated in the 35S::PdGA20ox1 stem tissues, suggesting a growth versus defense trade-off. Taken together, our results suggest that PdGA20ox1 functions to promote stem growth and wood formation in poplar, probably by activating GA signaling while coincidentally depressing defense signaling.

Phenotypic plasticity of sun and shade ecotypes of Stellaria longipes in response to light quality signaling, gibberellins and auxin.

Stellaria longipes plant communities (ecotypes) occur in several environmentally distinct habitats along the eastern slopes of southern Alberta's Rocky Mountains. One ecotype occurs in a prairie habitat at ∼1000 m elevation where Stellaria plants grow in an environment in which the light is filtered by taller neighbouring vegetation, i.e. sunlight with a low red to far-red (R/FR) ratio. This ecotype exhibits a high degree of phenotypic plasticity by increasing stem elongation in response to the low R/FR ratio light signal. Another Stellaria ecotype occurs nearby at ∼2400 m elevation in a much cooler alpine habitat, one where plants rarely experience low R/FR ratio shade light. Stem elongation of plants is largely regulated by gibberellins (GAs) and auxin, indole-3-acetic acid (IAA). Shoots of the prairie ecotype plants show increased IAA levels under low R/FR ratio light and they also increase their stem growth in response to applied IAA. The alpine ecotype plants show neither response. Plants from both ecotypes produce high levels of growth-active GA1 under low R/FR ratio light, though they differ appreciably in their catabolism of GA1. The alpine ecotype plants exhibit very high levels of GA8, the inactive product of GA1 metabolism, under both normal and low R/FR ratio light. Alpine origin plants may de-activate GA1 by conversion to GA8 via a constitutively high level of expression of the GA2ox gene, thereby maintaining their dwarf phenotype and exhibiting a reduced phenotypic plasticity in terms of shoot elongation. In contrast, prairie plants exhibit a high degree of phenotypic plasticity, using low R/FR ratio light-mediated changes in GA and IAA concentrations to increase shoot elongation, thereby accessing direct sunlight to optimize photosynthesis. There thus appear to be complex adaptation strategies for the two ecotypes, ones which involve modifications in the homeostasis of endogenous hormones.

Stress-related hormones and glycinebetaine interplay in protection of photosynthesis under abiotic stress conditions.

Plants subjected to abiotic stresses such as extreme high and low temperatures, drought or salinity, often exhibit decreased vegetative growth and reduced reproductive capabilities. This is often associated with decreased photosynthesis via an increase in photoinhibition, and accompanied by rapid changes in endogenous levels of stress-related hormones such as abscisic acid (ABA), salicylic acid (SA) and ethylene. However, certain plant species and/or genotypes exhibit greater tolerance to abiotic stress because they are capable of accumulating endogenous levels of the zwitterionic osmolyte-glycinebetaine (GB). The accumulation of GB via natural production, exogenous application or genetic engineering, enhances plant osmoregulation and thus increases abiotic stress tolerance. The final steps of GB biosynthesis occur in chloroplasts where GB has been shown to play a key role in increasing the protection of soluble stromal and lumenal enzymes, lipids and proteins, of the photosynthetic apparatus. In addition, we suggest that the stress-induced GB biosynthesis pathway may well serve as an additional or alternative biochemical sink, one which consumes excess photosynthesis-generated electrons, thus protecting photosynthetic apparatus from overreduction. Glycinebetaine biosynthesis in chloroplasts is up-regulated by increases in endogenous ABA or SA levels. In this review, we propose and discuss a model describing the close interaction and synergistic physiological effects of GB and ABA in the process of cold acclimation of higher plants.

Enhancing crop yield with the use of N-based fertilizers co-applied with plant hormones or growth regulators.

Crop yield, vegetative or reproductive, depends on access to an adequate supply of essential mineral nutrients. At the same time, a crop plant's growth and development, and thus yield, also depend on in situ production of plant hormones. Thus optimizing mineral nutrition and providing supplemental hormones are two mechanisms for gaining appreciable yield increases. Optimizing the mineral nutrient supply is a common and accepted agricultural practice, but the co-application of nitrogen-based fertilizers with plant hormones or plant growth regulators is relatively uncommon. Our review discusses possible uses of plant hormones (gibberellins, auxins, cytokinins, abscisic acid and ethylene) and specific growth regulators (glycine betaine and polyamines) to enhance and optimize crop yield when co-applied with nitrogen-based fertilizers. We conclude that use of growth-active gibberellins, together with a nitrogen-based fertilizer, can result in appreciable and significant additive increases in shoot dry biomass of crops, including forage crops growing under low-temperature conditions. There may also be a potential for use of an auxin or cytokinin, together with a nitrogen-based fertilizer, for obtaining additive increases in dry shoot biomass and/or reproductive yield. Further research, though, is needed to determine the potential of co-application of nitrogen-based fertilizers with abscisic acid, ethylene and other growth regulators.

Plant hormone-assisted early family selection in Pinus densiflora via a retrospective approach.

In an even-aged pine forest trees can vary considerably in stem size. We examined the basis for this anomaly using a retrospective approach. Twelve open-pollinated families of Pinus densiflora Sieb. et Zucc. were deliberately chosen for their variation in stem volumes at age 32 years. Seedlings obtained from these families were grown to age 6 months under optimal nursery conditions. Endogenous levels of growth hormones (auxin [IAA] and gibberellins [GAs]) and expression of the GA biosynthesis gene, PdGA20ox1, all assessed at age 3 months, were significantly correlated, across family, with seedling stem and/or shoot dry biomass at age 6 months. Retrospective comparisons of seedling growth, seedling stem tissue GA(20) and seedling stem expression levels of PdGA20ox1 were then made, across family, with tree stem growth at age 32 years. Age 6 months length and shoot dry biomass at age 6 months showed positive and significant Pearson's correlations with age 32 years tree stem diameters and a tree stem volume index, as did seedling stem tissue GA(20). Even seedling stem PdGA20ox1 expression levels were positively and near significantly (P = 0.088) correlated with age 32 years tree stem diameters. Auxin and GAs control nursery growth of seedlings at the family level, and this control also extends, for GAs at least, to field growth of older trees. We propose that family differences in PdGA20ox1 gene expression, and thus endogenous GA levels, may explain much of the natural variation seen for tree stem size in even-aged pine forests. If our hypothesis is correct, then the heritable components of variation in tree stem growth capacity should be predictable by hormonal and gene expression profiling. Such profiling, combined with the measurement of seedling phenotypic growth characters, could have the potential to accelerate the early selection of those conifer families that possess traits for inherently rapid stem wood growth.

A potential role for endogenous microflora in dormancy release, cytokinin metabolism and the response to fluridone in Lolium rigidum seeds.

BACKGROUND AND AIMS: Dormancy in Lolium rigidum (annual ryegrass) seeds can be alleviated by warm stratification in the dark or by application of fluridone, an inhibitor of plant abscisic acid (ABA) biosynthesis via phytoene desaturase. However, germination and absolute ABA concentration are not particularly strongly correlated. The aim of this study was to determine if cytokinins of both plant and bacterial origin are involved in mediating dormancy status and in the response to fluridone. METHODS: Seeds with normal or greatly decreased (by dry heat pre-treatment) bacterial populations were stratified in the light or dark and in the presence or absence of fluridone in order to modify their dormancy status. Germination was assessed and seed cytokinin concentration and composition were measured in embryo-containing or embryo-free seed portions. KEY RESULTS: Seeds lacking bacteria were no longer able to lose dormancy in the dark unless supplied with exogenous gibberellin or fluridone. Although these seeds showed a dramatic switch from active cytokinin free bases to O-glucosylated storage forms, the concentrations of individual cytokinin species were only weakly correlated to dormancy status. However, cytokinins of apparently bacterial origin were affected by fluridone and light treatment of the seeds. CONCLUSIONS: It is probable that resident microflora contribute to dormancy status in L. rigidum seeds via a complex interaction between hormones of both plant and bacterial origin. This interaction needs to be taken into account in studies on endogenous seed hormones or the response of seeds to plant growth regulators.

Light signaling and the phytohormonal regulation of shoot growth.

Shoot growth of dicot plants is rigorously controlled by the interactions of environmental cues with several groups of phytohormones. The signaling effects of light on shoot growth are of special interest, as both light irradiance and light quality change rapidly throughout the day, causing profound changes in stem elongation and leaf area growth. Among the several dicot species examined, we have focused on sunflower (Helianthus annuus L.) because its shoots are robust and their growth is highly plastic. Sunflower shoots thus constitute an ideal tissue for assessing responses to both light irradiance and light quality signals. Herein, we discuss the possible roles of gibberellins, auxin, ethylene, cytokinins and brassinosteroids in mediating the stem elongation and leaf area growth that is induced by shade light. To do this we uncoupled the plant's responses to changes in the red to far-red [R/FR] light ratio from its responses to changes in irradiance of photosynthetically active radiation [PAR]. Reducing each of R/FR light ratio and PAR irradiance results in increased sunflower stem elongation. However, the plant's response for leaf area growth differs considerably, with a low R/FR ratio generally promoting leaf area growth, whereas low irradiance PAR inhibits it. The increased stem elongation that occurs in response to lowering R/FR ratio and PAR irradiance is accomplished at the expense of leaf area growth. In effect, the low PAR irradiance signal overrides the low R/FR ratio signal in shade light's control of leaf growth and development. Three hormone groups, gibberellins, auxin and ethylene are directly involved in regulating these light-mediated shoot growth changes. Gibberellins and auxin function as growth promoters, with auxin likely acting as an up-regulator of gibberellin biosynthesis. Ethylene functions as a growth-inhibitor and probably interacts with gibberellins in regulating both stem and leaf growth of the sunflower shoot.

Potential for increased photosynthetic performance and crop productivity in response to climate change: role of CBFs and gibberellic acid.

We propose that targeting the enhanced photosynthetic performance associated with the cold acclimation of winter cultivars of rye (Secale cereale L.), wheat (Triticum aestivum L.), and Brassica napus L. may provide a novel approach to improve crop productivity under abiotic as well as biotic stress conditions. In support of this hypothesis, we provide the physiological, biochemical, and molecular evidence that the dwarf phenotype induced by cold acclimation is coupled to significant enhancement in photosynthetic performance, resistance to photoinhibition, and a decreased dependence on photoprotection through non-photochemical quenching which result in enhanced biomass production and ultimately increased seed yield. These system-wide changes at the levels of phenotype, physiology, and biochemistry appear to be governed by the family of C-repeat/dehydration-responsive family of transcription factors (CBF/DREB1). We relate this phenomenon to the semi-dwarf, gibberellic acid insensitive (GAI), cereal varieties developed during the "green revolution" of the early 1960s and 1970s. We suggest that genetic manipulation of the family of C-repeat/dehydration-responsive element binding transcription factors (CBF/DREB1) may provide a novel approach for the maintenance and perhaps even the enhancement of plant productivity under conditions of sub-optimal growth conditions predicted for our future climate.

Improving ryegrass-clover pasture dry matter yield and urea efficiency with gibberellic acid.

BACKGROUND: The effects of spraying gibberellic acid (GA3) at 20 or 30 g ha(-1), with or without application of urea, on pasture dry matter (DM) yield, herbage nitrogen (N) concentration and feed quality were investigated in 2011 and 2012 for managed pastoral systems in New Zealand across a range of sites, in both autumn and spring. RESULTS: On the Waikato site (autumn and spring, 2012), and at all five sites in 2011, liquid urea applied with GA3 at 20 or 30 g ha(-1) consistently produced significantly higher pasture shoot DM yield, relative to liquid urea alone. Application of GA3 alone reduced feed quality by lowering metabolizable energy, crude protein and organic matter digestibility values. However, a reduced feed quality was not observed when GA3 was applied together with liquid urea. Liquid urea applied with GA3 also reduced total N and nitrate-N concentration in herbage, relative to liquid urea applied alone. CONCLUSION: Application of GA3 together with liquid urea provides an opportunity for the strategic use of urea to meet both production and environmental goals.

Phytohormonal basis for the plant growth promoting action of naturally occurring biostimulators.

There is increasing interest in the use of naturally occurring 'biostimulators' for enhancing the growth of agricultural and horticultural crops. Bacteria, fungi and protozoa, as well as marine algae-based seaweed extracts, can produce or contain biostimulators. The activity of biostimulators to promote plant growth is often attributed to their ability to directly or indirectly provide mineral nutrients (mostly N, but also P, S and other macro- and micro-nutrients) to plants. Alternatively, biostimulators are postulated to increase the plant's ability to assimilate these mineral nutrients, often in return for photo-assimilates (as occurs with certain bacteria and fungi associations). Although optimal growth of plants depends on the availability of adequate mineral nutritients, that growth (and also development, including reproduction) is also regulated by plant hormones (phytohormones), including gibberellins, auxins and cytokinins. This review describes and discusses the evidence that the presence or application of biostimulators also increases plant growth directly via phytohormone action and also influences the plant's ability to control its own hormone biosynthesis and homeostasis. Finally, it discusses the need for a better understanding of the role(s) that are played by the naturally occurring biostimulators associated with the plant in the crop field. It is suggested that better understanding will allow for optimal crop yield returns, since disruptions of phytohormone homeostasis in plant organs and tissues can yield either beneficial or sub-optimal outcomes.

Gibberellin 3-oxidase gene expression patterns influence gibberellin biosynthesis, growth, and development in pea.

Gibberellins (GAs) are key modulators of plant growth and development. PsGA3ox1 (LE) encodes a GA 3ß-hydroxylase that catalyzes the conversion of GA20 to biologically active GA1. To further clarify the role of GA3ox expression during pea (Pisum sativum) plant growth and development, we generated transgenic pea lines (in a lele background) with cauliflower mosaic virus-35S-driven expression of PsGA3ox1 (LE). PsGA3ox1 transgene expression led to higher GA1 concentrations in a tissue-specific and development-specific manner, altering GA biosynthesis and catabolism gene expression and plant phenotype. PsGA3ox1 transgenic plants had longer internodes, tendrils, and fruits, larger stipules, and displayed delayed flowering, increased apical meristem life, and altered vascular development relative to the null controls. Transgenic PsGA3ox1 overexpression lines were then compared with lines where endogenous PsGA3ox1 (LE) was introduced, by a series of backcrosses, into the same genetic background (BC LEle). Most notably, the BC LEle plants had substantially longer internodes containing much greater GA1 levels than the transgenic PsGA3ox1 plants. Induction of expression of the GA deactivation gene PsGA2ox1 appears to make an important contribution to limiting the increase of internode GA1 to modest levels for the transgenic lines. In contrast, PsGA3ox1 (LE) expression driven by its endogenous promoter was coordinated within the internode tissue to avoid feed-forward regulation of PsGA2ox1, resulting in much greater GA1 accumulation. These studies further our fundamental understanding of the regulation of GA biosynthesis and catabolism at the tissue and organ level and demonstrate that the timing/localization of GA3ox expression within an organ affects both GA homeostasis and GA1 levels, and thereby growth.

Role of CBFs as integrators of chloroplast redox, phytochrome and plant hormone signaling during cold acclimation.

Cold acclimation of winter cereals and other winter hardy species is a prerequisite to increase subsequent freezing tolerance. Low temperatures upregulate the expression of C-repeat/dehydration-responsive element binding transcription factors (CBF/DREB1) which in turn induce the expression of COLD-REGULATED (COR) genes. We summarize evidence which indicates that the integration of these interactions is responsible for the dwarf phenotype and enhanced photosynthetic performance associated with cold-acclimated and CBF-overexpressing plants. Plants overexpressing CBFs but grown at warm temperatures mimic the cold-tolerant, dwarf, compact phenotype; increased photosynthetic performance; and biomass accumulation typically associated with cold-acclimated plants. In this review, we propose a model whereby the cold acclimation signal is perceived by plants through an integration of low temperature and changes in light intensity, as well as changes in light quality. Such integration leads to the activation of the CBF-regulon and subsequent upregulation of COR gene and GA 2-oxidase (GA2ox) expression which results in a dwarf phenotype coupled with increased freezing tolerance and enhanced photosynthetic performance. We conclude that, due to their photoautotrophic nature, plants do not rely on a single low temperature sensor, but integrate changes in light intensity, light quality, and membrane viscosity in order to establish the cold-acclimated state. CBFs appear to act as master regulators of these interconnecting sensing/signaling pathways.

Impact of eastern dwarf mistletoe (Arceuthobium pusillum) on host white spruce (Picea glauca) development, growth and performance across multiple scales.

Infection by eastern dwarf mistletoe (Arceuthobium pusillum) modifies needle and branch morphology and hastens white spruce (Picea glauca) mortality. We examined potential causal mechanisms and assessed the impacts of infection-induced alterations to host development and performance across scales ranging from needle hormone contents to bole expansion. Needles on infected branches (IBs) possessed higher total cytokinin (CK) and lower abscisic acid contents than needles on uninfected branches (UBs). IBs exhibited greater xylem growth than same-aged UBs, which is consistent with the promotive effect of CKs on vascular differentiation and organ sink strength. Elevated CK content may also explain the dense secondary and tertiary branching observed at the site of infection, i.e. the formation of 'witches' brooms' with significantly lower light capture efficiencies. Observed hormone perturbations were consistent with higher rates of transpiration, lower water use efficiencies (WUEs) and more negative needle carbon isotope ratios observed for IBs. Observed reductions in needle size allowed IBs to compensate for reduced hydraulic conductivity. Severe infections resulted in dramatically decreased diameter growth of the bole. It seems likely that the modifications to host hormone contents by eastern dwarf mistletoe infection led white spruce trees to dedicate a disproportionate fraction of their photoassimilate and other resources to self-shaded branches with low WUE. This would have decreased the potential for fixed carbon accumulation, generating a decline in the whole-tree resource pool. As mistletoe infections grew in size and the number of IBs increased, this burden was manifested as increasingly greater reductions in bole growth.

Ethylene involvement in silique and seed development of canola, Brassica napus L.

A wide range of plant hormones, including gibberellins (GAs) and auxins are known to be involved in regulating seed and fruit growth and development. Changes in ethylene biosynthesis are also associated with seed and fruit development, but ethylene's role in these processes is poorly understood, as is its possible interaction with the other plant hormones. A major complication of investigating ethylene-induced regulation of developmental processes is ethylene's biphasic mode of action. To investigate ethylene's actions and interactions we used a 1-amino-cyclopropane-1-carboxylic acid (ACC) deaminase transgenic canola line. This line evolves significantly less ethylene from its siliques and seeds, relative to plants from a wild type (WT) background. Plants of the transgenic line also had smaller siliques which were associated with reductions in both seed size and seed number. Application of ethephon, a compound that produces ethylene, to plants of the transgenic line restored the WT phenotype for both siliques and seeds. Application of the same dose of ethephon to WT plants diminished both silique and seed development, showing ethylene's biphasic effect and effectively producing the ACC deaminase transgenic phenotype. There were significant decreases in endogenous concentrations of GA(1) and GA(4) and also of indole-3-acetic acid (IAA), between WT seeds and seedless siliques and seeds and siliques from the transgenic line plants. These differences were emphasized during early stages (10-20 days after pollination) of seed and silique development. The above results strongly suggest that ethylene interacts with other endogenous plant hormones in regulating silique and seed development and growth in WT lines of canola.

Flowering in snow tussock (Chionochloa spp.) is influenced by temperature and hormonal cues.

Snow tussocks (Chionochloa spp.) in New Zealand exhibit extreme mast (episodic) seeding which has important implications for plant ecology and plant-insect interactions. Heavy flowering appears to be triggered by very warm/dry summers in the preceding year. In order to investigate the physiological basis for mast flowering, mature snow tussock plants in the field and younger plants in a glasshouse and shadehouse were subjected to a range of manipulative treatments. Field treatments included combinations of warming, root pruning and applications of two native gibberellins (GAs) GA3, which is known to be highly floral inductive and GA4, which is associated with continued floral apex development in another long-day grass. Warming, GA3 alone and especially warming+GA3, significantly promoted flowering, as did applications of GA4 alone and GA4+CCC (2-chloroethyltrimethylammonium chloride, which is a known synergist of GA3-induced flowering in the annual grass, Lolium temulentum L.). Our results provide support for the concept that mast flowering events in tussock species are causally related to high temperature-induced increases in endogenous gibberellin levels. It is likely that GAs (endogenous or applied) promote the continued development of a previously long-day induced floral apex. In addition to the promotion of flowering, applied GA3 also disturbed the plant's innate resource threshold requirements, as shown by the death, over winter, of many non-flowering tillers. Applied GA4 did not show this effect, likely due to its rapid catabolic metabolism to an inactive form. High temperature-induced flowering mediated by elevated levels of endogenous floral-promotive GAs could have important implications for regulating the evolutionary interaction between these masting plants and their seed predators.

Tissue-specific regulation of gibberellin biosynthesis in developing pea seeds.

Previous work suggests that gibberellins (GAs) play an important role in early seed development. To more fully understand the roles of GAs throughout seed development, tissue-specific transcription profiles of GA metabolism genes and quantitative profiles of key GAs were determined in pea (Pisum sativum) seeds during the seed-filling development period (8-20 d after anthesis [DAA]). These profiles were correlated with seed photoassimilate acquisition and storage as well as morphological development. Seed coat growth (8-12 DAA) and the subsequent dramatic expansion of branched parenchyma cells were correlated with both transcript abundance of GA biosynthesis genes and the concentration of the growth effector GA, GA(1). These results suggest GA(1) involvement in determining the rate of seed coat growth and sink strength. The endosperm's PsGA20ox transcript abundance and the concentration of GA(20) increased markedly as the endosperm reached its maximum volume (12 DAA), thus providing ample GA(20) substrate for the GA 3-oxidases present in both the embryo and seed coat. Furthermore, PsGA3ox transcript profiles and trends in GA(1) levels in embryos at 10 to 16 DAA and also in embryo axes at 18 DAA suggest localized GA(1)-induced growth in these tissues. A shift from synthesis of GA(1) to that of GA(8) occurred after 18 DAA in the embryo axis, suggesting that deactivation of GA(1) to GA(8) is a likely mechanism to limit embryo axis growth and allow embryo maturation to proceed. We hypothesize that GA biosynthesis and catabolism are tightly regulated to bring about the unique developmental events that occur during seed growth, development, and maturation.