Physiological Impacts of Elevated CO2 Concentration Ranging from Molecular to Whole Plant Responses

The dynamics of the terrestrial ecosystems depend on interactions between a number of biogeochemical cycles (i.e. carbon, nutrient, and hydrological cycles) that may be modified by human actions. Conversely, terrestrial ecosystems are important components of these cycles that create the sources and sinks of important greenhouse gases (e.g. carbon dioxide, methane, nitrous oxide). Especially, carbon is exchanged naturally among these ecosystems and the atmosphere through photosynthesis, respiration, decomposition, and combustion processes. Continuous increase of atmospheric carbon dioxide (CO₂) concentration has led to extensive research over the last two decades, during which more then 1 400 scientific papers describing impacts of elevated [CO₂] (EC) on photosynthesis have been published. However, the degree of response is very variable, depending on species, growing conditions, mineral nutrition, and duration of CO₂ enrichment. In this review, I have summarised the major physiological responses of plants, in particular of trees, to EC including molecular and primary, especially photosynthetic, physiological responses. Likewise, secondary (photosynthate translocation and plant water status) and tertiary whole plant responses including also plant to plant competition are shown.     

Effect of Elevated CO2 on Photosynthesis and Chlorophyll Fluorescence of Rose Plants Grown at High Temperature and High Photosynthetic Photon Flux Density

Gas exchange and chlorophyll (Chl) fluorescence were measured on young mature leaves of rose plants (Rosa hybrida cvs. First Red and Twingo) grown in two near-to-tight greenhouses, one under control ambient CO₂ concentration, AC (355 µmol mol^–1) and one under CO₂ enrichment, EC (700 µmol mol^–1), during four flushes from late June to early November. Supply of water and mineral elements was non-limiting while temperature was allowed to rise freely during daytime. Leaf diffusive conductance was not significantly reduced at EC but net photosynthetic rate increased by more than 100 %. Although the concentration of total non-structural saccharides was substantially higher in the leaves from the greenhouse with EC, _PS2 (quantum efficiency of radiation use) around noon was not significantly reduced at EC indicating that there was no down-regulation of electron transport. Moreover, CO₂ enrichment did not cause any increase in the risk of photo-damage, as estimated by the 1 – q_P parameter. Non-photochemical quenching was even higher in the greenhouse with EC during the two summer flushes, when temperature and photosynthetic photon flux density (PPFD) were the highest. Hence rose photosynthesis benefits strongly from high concentrations of atmospheric CO₂ at both high and moderate temperatures and PPFD.     

Different Responses of Norway Spruce Needles from Shaded and Exposed Crown Layers to the Prolonged Exposure to Elevated CO2 Studied by Various Chlorophyll a Fluorescence Techniques

The acclimation depression of capacity of photon utilisation in photochemical reactions of photosystem 2 (PS2) can develop already after three months of cultivation of the Norway spruces (Picea abies [L.] Karst.) under elevated concentrations of CO₂ (i.e., ambient, AC, + 350 µmol(CO₂) mol^–1 = EC) in glass domes with adjustable windows. To examine the role that duration of EC plays in acclimation response, we determined pigment contents, rate of photosynthesis, and parameters of chlorophyll a fluorescence for sun and shade needles after three seasons of EC exposure. We found responses of shaded and exposed needles to EC. Whereas the shaded needles still profited from the EC and revealed stimulated electron transport, for the exposed needles the stimulation of both electron transport activity and irradiance saturated rate of CO₂ assimilation (P _Nmax) under EC already disappeared. No signs of the PS2 impairment were observed as judged from high values of potential quantum yield of PS2 photochemistry (F_V/F_M) and uniform kinetics of Q_A reoxidation for all variants. Therefore, the long-term acclimation of the sun-exposed needles to EC is not necessarily accompanied with the damage to the PS2 reaction centres. The eco-physiological significance of the reported differentiation between the responses of shaded and sun exposed needles to prolonged EC may be in changed contribution of the upper and lower crown layers to the production activity of the tree. Whereas for the AC spruces, P _Nmax of shaded needles was only less than 25 % compared to exposed ones, for the EC spruces the P _Nmax of shaded needles reached nearly 40 % of that estimated for the exposed ones. Thus, the lower shaded part of the crown may become an effective consumer of CO₂.     

A regulatory link between ER-associated protein degradation and the unfolded-protein response

Ubiquitin conjugation during endoplasmic-reticulum-associated degradation (ERAD) depends on the activity of Ubc7. Here we show that Ubc1 acts as a further ubiquitin-conjugating enzyme in this pathway. Absence of both enzymes results in marked stabilization of an ERAD substrate and induction of the unfolded-protein response (UPR). Furthermore, basic ERAD activity is sufficient to eliminate unfolded proteins under normal conditions. However, when stress is applied, the UPR is required to increase ERAD activity. We thus demonstrate, for the first time, a regulatory loop between ERAD and the UPR, which is essential for normal growth of yeast cells.     

Differential role of neurokinin receptors in human lymphocyte and monocyte chemotaxis

The precise nature of neurokin receptor involvement in human immune cell chemotaxis is unclear. This study therefore sought to directly compare the chemotactic effects of neurokinins on human T lymphocytes and monocytes. Substance P was found to have a similar dose-dependent chemotactic action on T lymphocyte and monocyte populations. In contrast, T lymphocytes were found to be more responsive than monocytes both to the highly selective NK-1 agonist, [Sar(9)Met O(2)(11)]-substance P, and also to the NK-2 selective agonist, beta-alanine neurokinin A((4-10)). Consistent with these findings, substance P-induced chemotaxis of both T lymphocyte and monocytes was attenuated by the selective NK-1 antagonist LY303870. However, the selective NK-2 antagonist MEN 10,376 was only effective in inhibiting the T lymphocyte response. The study confirms that neurokinins have chemotactic actions on immune cells and indicates important functional differences between human T lymphocyte and monocyte responses. This provides a potential mechanism by which the nervous system can selectively influence cellular recruitment in inflammatory disease.     

Sex-specific Trans-regulatory Variation on the Drosophila melanogaster X Chromosome

The X chromosome constitutes a unique genomic environment because it is present in one copy in males, but two copies in females. This simple fact has motivated several theoretical predictions with respect to how standing genetic variation on the X chromosome should differ from the autosomes. Unmasked expression of deleterious mutations in males and a lower census size are expected to reduce variation, while allelic variants with sexually antagonistic effects, and potentially those with a sex-specific effect, could accumulate on the X chromosome and contribute to increased genetic variation. In addition, incomplete dosage compensation of the X chromosome could potentially dampen the male-specific effects of random mutations, and promote the accumulation of X-linked alleles with sexually dimorphic phenotypic effects. Here we test both the amount and the type of genetic variation on the X chromosome within a population of Drosophila melanogaster, by comparing the proportion of X linked and autosomal trans-regulatory SNPs with a sexually concordant and discordant effect on gene expression. We find that the X chromosome is depleted for SNPs with a sexually concordant effect, but hosts comparatively more SNPs with a sexually discordant effect. Interestingly, the contrasting results for SNPs with sexually concordant and discordant effects are driven by SNPs with a larger influence on expression in females than expression in males. Furthermore, the distribution of these SNPs is shifted towards regions where dosage compensation is predicted to be less complete. These results suggest that intrinsic properties of dosage compensation influence either the accumulation of different types of trans-factors and/or their propensity to accumulate mutations. Our findings document a potential mechanistic basis for sex-specific genetic variation, and identify the X as a reservoir for sexually dimorphic phenotypic variation. These results have general implications for X chromosome evolution, as well as the genetic basis of sex-specific evolutionary change.     

Endothelial cell and podocyte autophagy synergistically protect from diabetes-induced glomerulosclerosis

The glomerulus is a highly specialized capillary tuft, which under pressure filters large amounts of water and small solutes into the urinary space, while retaining albumin and large proteins. The glomerular filtration barrier (GFB) is a highly specialized filtration interface between blood and urine that is highly permeable to small and midsized solutes in plasma but relatively impermeable to macromolecules such as albumin. The integrity of the GFB is maintained by molecular interplay between its 3 layers: the glomerular endothelium, the glomerular basement membrane and podocytes, which are highly specialized postmitotic pericytes forming the outer part of the GFB. Abnormalities of glomerular ultrafiltration lead to the loss of proteins in urine and progressive renal insufficiency, underlining the importance of the GFB. Indeed, albuminuria is strongly predictive of the course of chronic nephropathies especially that of diabetic nephropathy (DN), a leading cause of renal insufficiency. We found that high glucose concentrations promote autophagy flux in podocyte cultures and that the abundance of LC3B II in podocytes is high in diabetic mice. Deletion of Atg5 specifically in podocytes resulted in accelerated diabetes-induced podocytopathy with a leaky GFB and glomerulosclerosis. Strikingly, genetic alteration of autophagy on the other side of the GFB involving the endothelial-specific deletion of Atg5 also resulted in capillary rarefaction and accelerated DN. Thus autophagy is a key protective mechanism on both cellular layers of the GFB suggesting autophagy as a promising new therapeutic strategy for DN.