Association between tree-ring and needle δ<Superscript>13</Superscript>C and leaf gas exchange in <Emphasis Type="Italic">Pinus halepensis</Emphasis> under semi-arid conditions

Associations between δ¹³C values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO₂ uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of δ¹³C fractionation in C₃ plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated δ¹³C values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and δ¹³C responses to changing environmental conditions. Seasonal cycles of growth and δ¹³C (up to ~4‰) significantly correlated (P<0.01) with photosynthetically active radiation, vapour pressure deficit, air temperature, and soil water content. The irrigation significantly increased leaf net assimilation, stomatal conductance and needle and tree-ring growth rate, and markedly decreased needle and tree-ring δ¹³C values and its sensitivity to environmental parameters. The δ¹³C estimates derived from gas-exchange parameters, and weighted by assimilation, compared closely with seasonal and inter-annual δ¹³C values of needle- and tree-ring tissue. Higher stomatal conductances of the irrigated trees (0.22 vs. 0.08 mol m⁻² s⁻¹ on average) corresponded with ~2.0‰ lower average δ¹³C values, both measured and derived. Derived and measured δ¹³C values also indicated that needle growth, which occurs throughout the stressful summer was supported by carbon from concurrent, low rate assimilation. For Aleppo pine under semi-arid and irrigated conditions, the δ¹³C of tree-ring and needle material proved, in general, to be a reasonable indicator of integrated leaf gas-exchange properties.     

Tissue-specific expression of Populus C19 GA 2-oxidases differentially regulate above- and below-ground biomass growth through control of bioactive GA concentrations

? Here, we studied the poplar C(19) gibberellin 2-oxidase (GA2ox) gene subfamily. We show that a set of paralogous gene pairs differentially regulate shoot and root development. ? PtGA2ox4 and its paralogous gene PtGA2ox5 are primarily expressed in aerial organs, and overexpression of PtGA2ox5 produced a strong dwarfing phenotype characteristic of GA deficiency. Suppression of PtGA2ox4 and PtGA2ox5 led to increased biomass growth, but had no effect on root development. By contrast, the PtGA2ox2 and PtGA2ox7 paralogous pair was predominantly expressed in roots, and when these two genes were RNAi-suppressed it led to a decrease of root biomass. ? The morphological changes in the transgenic plants were underpinned by tissue-specific increases in bioactive GAs that corresponded to the predominant native expression of the targeted paralogous gene pair. Although RNAi suppression of both paralogous pairs led to changes in wood development, they were much greater in the transgenics with suppressed PtGA2ox4 and PtGA2ox5. The degree of gene suppression in independent events was strongly associated with phenotypes, demonstrating dose-dependent control of growth by GA2ox RNA concentrations. ? The expression and transgenic modifications reported here show that shoot- and leaf-expressed PtGA2ox4 and PtGA2ox5 specifically restrain aerial shoot growth, while root-expressed PtGA2ox2 and PtGA2ox7 promote root development.     

Large daily variation in 13C-enrichment of leaf-respired CO2 in two Quercus forest canopies

The use of the 13C : 12C isotopic ratio (delta13C) of leaf-respired CO2 to trace carbon fluxes in plants and ecosystems is limited by little information on temporal variations in delta13C of leaf dark-respired CO2 (delta13Cr) under field conditions. Here, we explored variability in delta13Cr and its relationship to key respiratory substrates from collections of leaf dark-respired CO2, carbohydrate extractions and gas exchange measurements over 24-h periods in two Quercus canopies. Throughout both canopies, delta13Cr became progressively 13C-enriched during the photoperiod, by up to 7%, then 13C-depleted at night relative to the photoperiod. This cycle could not be reconciled with delta13C of soluble sugars (delta13Css), starch (delta13Cst), lipids (delta13Cl), cellulose (delta13Cc) or with calculated photosynthetic discrimination (Delta). However, photoperiod progressive enrichment in delta13Cr was correlated with cumulative carbon assimilation (r2 = 0.91). We concluded that there is considerable short-term variation in delta13Cr in forest canopies, that it is consistent with current hypotheses for 13C fractionation during leaf respiration, that leaf carbohydrates cannot be used as surrogates for delta13Cr, and that diel changes in leaf carbohydrate status could be used to predict changes in delta13Cr empirically.     

Physiology-phenology interactions in a productive semi-arid pine forest

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D(L)) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g(s)) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g(s )sensitivity to D(L) below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis-phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.     

An assessment of the relationship between chlorophyll a fluorescence and CO2 gas exchange from field measurements on a moss and lichen

The relationship between CO₂ exchange and relative electron-transport rate through photosystem II (ETR, measured using chlorophyll a fluorescence) was determined for a moss and a green algal lichen, photobiont probably Trebouxia sp., in the field in Antarctica. Net photosynthesis (NP) and dark respiration (DR) were measured over temperatures from zero to 25 °C and gross photosynthesis (GP) calculated (GP = NP + DR). The strong response of DR to temperature in these organisms resulted in substantial changes in CO₂ exchange rates. The moss Bryum argenteum Hedw. showed a strong, linear relationship between GP and ETR. This was an unexpected result since mosses are C₃ plants and, in higher plants, this group normally has a curvilinear GP versus ETR relationship. It is suggested that suppression of DR in the light might be involved. The lichen, Umbilicaria aprina Nyl., had nonlinear relationships between ETR and GP that were different at each measurement temperature. In some cases the lowest ETR was at the higher CO₂ exchange rates. It is suggested that these relationships are the result of strong quenching mechanisms that are inversely proportional to GP. The results support a growing impression that the relationships between ETR and CO₂ exchange are complex in these organisms and different from those found for higher plants. Received: 24 November 1997 / Accepted 2 May 1998     

Repression of gibberellin biosynthesis or signaling produces striking alterations in poplar growth,morphology, and flowering

We modified gibberellin (GA) metabolism and signaling in transgenic poplars using dominant transgenes and studied their effects for 3 years under field conditions. The transgenes that we employed either reduced the bioactive GAs, or attenuated their signaling. The majority of transgenic trees had significant and in many cases dramatic changes in height, crown architecture, foliage morphology, flowering onset, floral structure, and vegetative phenology. Most transgenes elicited various levels of height reduction consistent with the roles of GA in elongation growth. Several other growth traits were proportionally reduced, including branch length, internode distance, and leaf length. In contrast to elongation growth, stem diameter growth was much less affected, suggesting that semi-dwarf trees in dense stands might provide high levels of biomass production and carbon sequestration. The severity of phenotypic effects was strongly correlated with transgene expression among independent transgenic events, but often in a non-linear manner, the form of which varied widely among constructs. The majority of semi-dwarfed, transgenic plants showed delayed bud flush and early bud set, and expression of a native GAI transgene accelerated first time flowering in the field. All of the phenotypic changes observed in multiple years were stable over the 3 years of field study. Our results suggest that transgenic modification of GA action may be useful for producing semi-dwarf trees with modified growth and morphology for horticulture and other uses.     

Incorporating climate change into ecosystem service assessments and decisions: a review

Climate change is having a significant impact on ecosystem services and is likely to become increasingly important as this phenomenon intensifies. Future impacts can be difficult to assess as they often involve long timescales, dynamic systems with high uncertainties, and are typically confounded by other drivers of change. Despite a growing literature on climate change impacts on ecosystem services, no quantitative syntheses exist. Hence, we lack an overarching understanding of the impacts of climate change, how they are being assessed, and the extent to which other drivers, uncertainties, and decision making are incorporated. To address this, we systematically reviewed the peer‐reviewed literature that assesses climate change impacts on ecosystem services at subglobal scales. We found that the impact of climate change on most types of services was predominantly negative (59% negative, 24% mixed, 4% neutral, 13% positive), but varied across services, drivers, and assessment methods. Although uncertainty was usually incorporated, there were substantial gaps in the sources of uncertainty included, along with the methods used to incorporate them. We found that relatively few studies integrated decision making, and even fewer studies aimed to identify solutions that were robust to uncertainty. For management or policy to ensure the delivery of ecosystem services, integrated approaches that incorporate multiple drivers of change and account for multiple sources of uncertainty are needed. This is undoubtedly a challenging task, but ignoring these complexities can result in misleading assessments of the impacts of climate change, suboptimal management outcomes, and the inefficient allocation of resources for climate adaptation.     

Increase in water-use efficiency and underlying processes in pine forests across a precipitation gradient in the dry Mediterranean region over the past 30 years

Motivated by persistent predictions of warming and drying in the entire Mediterranean and other regions, we have examined the interactions of intrinsic water-use efficiency (W _i) with environmental conditions in Pinus halepensis. We used 30-year (1974–2003) tree-ring records of basal area increment (BAI) and cellulose ¹³C and ¹⁸O composition, complemented by short-term physiological measurements, from three sites across a precipitation (P) gradient (280–700 mm) in Israel. The results show a clear trend of increasing W _i in both the earlywood (EW) and latewood (LW) that varied in magnitude depending on site and season, with the increase ranging from ca. 5 to 20% over the study period. These W _i trends were better correlated with the increase in atmospheric CO₂ concentration, C _a, than with the local increase in temperature (~0.04°C year⁻¹), whereas age, height and density variations had minor effects on the long-term isotope record. There were no trends in P over time, but W _i from EW and BAI were dependent on the interannual variations in P. From reconstructed C _i values, we demonstrate that contrasting gas-exchange responses at opposing ends of the hydrologic gradient underlie the variation in W _i sensitivity to C _a between sites and seasons. Under the mild water limitations typical of the main seasonal growth period, regulation was directed at increasing C _i/C _a towards a homeostatic set-point observed at the most mesic site, with a decrease in the W _i response to C _i with increasing aridity. With more extreme drought stress, as seen in the late season at the drier sites, the response was W _i driven, and there was an increase in the W _i sensitivity to C _a with aridity and a decreasing sensitivity of C _i to C _a. The apparent C _a-driven increases in W _i can help to identify the adjustments to drying conditions that forest ecosystems can make in the face of predicted atmospheric change.     

Research resource: Haploinsufficiency of receptor activity-modifying protein-2 (RAMP2) causes reduced fertility, hyperprolactinemia, skeletal abnormalities, and endocrine dysfunction in mice

Receptor activity-modifying protein-2 (RAMP2) is a single-pass transmembrane protein that can regulate the trafficking, ligand binding, and signaling of several G protein-coupled receptors (GPCR). The most well-characterized role of RAMP2 is in the regulation of adrenomedullin (AM) binding to calcitonin receptor-like receptor (CLR), and our previous studies using knockout mouse models support this canonical signaling paradigm. For example, Ramp2(-/-) mice die at midgestation with a precise phenocopy of the AM(-/-) and Calcrl(-/-) mice. In contrast, Ramp2(+/-) mice are viable and exhibit an expanded variety of phenotypes that are distinct from those of Calcrl(+/-) mice. Using Ramp2(+/-) female mice, we demonstrate that a modest decrease in Ramp2 expression causes severe reproductive defects characterized by fetal growth restriction, fetal demise, and postnatal lethality that is independent of the genotype and gender of the offspring. Ramp2(+/-) female mice also exhibit hyperprolactinemia during pregnancy and in basal conditions. Consistent with hyperprolactinemia, Ramp2(+/-) female mice have enlarged pituitary glands, accelerated mammary gland development, and skeletal abnormalities including delayed bone development and decreased bone mineral density. Because RAMP2 has been shown to associate with numerous GPCR, it is likely that signaling of one or more of these GPCR is compromised in Ramp2(+/-) mice, yet the precise identification of these receptors remains to be elucidated. Taken together, this work reveals an essential role for RAMP2 in endocrine physiology and provides the first in vivo evidence for a physiological role of RAMP2 beyond that of AM/CLR signaling.     

SHORT INTERNODES-like genes regulate shoot growth and xylem proliferation in Populus

? Genes controlling plant growth and form are of considerable interest, because they affect survival and productivity traits, and are largely unknown or poorly characterized. The SHORT INTERNODES(SHI) gene is one of a 10-member SHI-RELATED SEQUENCE (SRS) gene family in Arabidopsis that includes important developmental regulators. ? Using comparative sequence analysis of the SRS gene families in poplar and Arabidopsis, we identified two poplar proteins that are most similar to SHI and its closely related gene STYLISH1 (STY1). The two poplar genes are very similar in sequence and expression and are therefore probably paralogs with redundant functions. ? RNAi suppression of the two Populus genes enhanced shoot and root growth, whereas the overexpression of Arabidopsis SHI in poplar reduced internode and petiole length. The suppression of the two genes increased fiber length and the proportion of xylem tissue, mainly through increased xylem cell proliferation. The transgenic modifications were also associated with significant changes in the concentrations of gibberellins and cytokinin. ? We conclude that Populus SHI-RELATED SEQUENCE (SRS) genes play an important role in the regulation of vegetative growth, including wood formation, and thus could be useful tools for the modification of biomass productivity, wood quality or plant form.