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The control of inorganic phosphate homeostasis is mediated through the activity of sodium-coupled Pi transporters located in the intestine, kidneys, and bone. To study these transporters in either the native tissue or after heterologous expression, it is very important to use specific inhibitors of the studied transporter, in order to know the corresponding relevance in the total Pi uptake and to differentiate from the activity of other transporters. Inhibitors are also necessary as drugs for treating Pi homeostasis disorders. Under normal physiological conditions, the renal and intestinal excretion of Pi matches dietary intestinal absorption, but when the number of non-functional nephrons increase in chronic kidney disease and end-stage renal disease, the excretion of surplus Pi is progressively impaired, thereby increasing the risk of hyperphosphatemia and Pi toxicity. When the compensatory mechanisms that increase Pi excretion fail, Pi toxicity can only be prevented by reducing the intestinal absorption of Pi through phosphate binders that reduced the free Pi concentration in the lumen, and inhibitors of intestinal Pi transporters and of the paracellular absorption route. Although many potentially interesting inhibitors have been reported to date, only a few are available for experimental purposes, and even fewer have been used in independent clinical trials. In this review, we summarize the different groups of compounds reported to date as inhibitors of Pi transport. To help understand and characterize the inhibition mechanisms, we also summarize the kinetic analysis approaches and screening methods that could be applied.

The cloning of the renal NaPi-2a (SLC34A1) and NaPi-2c (SLC34A3) phosphate transporters has made it possible to characterize the molecular and biophysical regulation of renal proximal tubular reabsorption of inorganic phosphate (Pi). Dietary factors, such as Pi and K, and several hormones and phosphatonins, including parathyroid hormone (PTH), fibroblast growth factor 23 (FGF23), and glucocorticoids, regulate the transporters through various transcriptional, translational, and post-translational mechanisms that involve acute trafficking via endocytosis or exocytosis, interactions with PDZ domain proteins, lipid microdomains, and diffusion and clustering in the apical brush border membrane. The visualization of these trafficking events by means of novel microscopy techniques that includes fluorescence lifetime imaging microscopy (FLIM), Förster resonance energy transfer (FRET), fluctuation correlation spectroscopy (FCS), and modulation tracking (MT), is the primary focus of this review.

The taxonomic status of the Pieris napi -complex and similar species which occur in China are revised. Relevant species distributed in the adjacent regions were included to clarify the status of Chinese species and were briefly revised. All those species are described and illustrated and new synonyms are established. A molecular phylogenetic analysis is also performed on the species group including similar species, to investigate the phylogenetic relationships between taxa. Species of the Pieris napi -complex that occur in China and adjacent regions are redefined, with four similar species excluded ( P. melaina , P. extensa , P. chumbiensis gyantsensis and P. melete ). A distribution map and keys of the complex including similar species are provided. The taxon P. mihon Yakovlev, 2006 stat. nov. is raised from subspecies to species status; P. narina Verity, 1908 stat. rev. is confirmed as a distinct species rather than a subspecies of P. ochsenheimeri ; Pieris euorientis Verity, 1908 stat. rev. is recovered as a distinct species sister to P. dulcinea . Two taxa, ssp. sauron and ssp. bryonides are moved from subspecies of P. euorientis and P. bryoniae , respectively, to P. napi , i.e. P. napi sauron Yakovlev, 2004 comb. nov and P. napi bryonides Sheljuzhko, 1910 comb. rev. A new synonym is proposed: Pieris ochsenheimeri tianshansis Tadokoro, Shinkawa & Wang, 2014, new synonym of P. mihon Yakovlev, 2006. A new mistaken identification is proposed: Pieris dulcinea kneitzi is a misidentification of Pieris erutae kneitzi Eitschberger, 1983 comb. rev. Five Chinese species belonging to the Pieris napi -complex were confirmed, namely P. narina , P. mihon , P. latouchei , P. dulcinea , and P. erutae . Among them, two species, P. mihon Yakovlev, 2006 and Pieris narina Verity, 1908, are newly recorded from China. The taxonomic status of Pieris steinigeri Eitschberger, 1983 and Pieris bryoniae sifanica Grum-Grshimailo, 1895 is also discussed.

Seasons impose different selection pressures on organisms through contrasting environmental conditions. How such seasonal evolutionary conflict is resolved in organisms whose lives span across seasons remains underexplored. Through field experiments, laboratory work, and citizen science data analyses, we investigate this question using two closely related butterflies ( Pieris rapae and P. napi ). Superficially, the two butterflies appear highly ecologically similar. Yet, the citizen science data reveal that their fitness is partitioned differently across seasons. Pieris rapae have higher population growth during the summer season but lower overwintering success than do P. napi . We show that these differences correspond to the physiology and behavior of the butterflies. Pieris rapae outperform P. napi at high temperatures in several growth season traits, reflected in microclimate choice by ovipositing wild females. Instead, P. rapae have higher winter mortality than do P. napi . We conclude that the difference in population dynamics between the two butterflies is driven by seasonal specialization, manifested as strategies that maximize gains during growth seasons and minimize harm during adverse seasons, respectively. Seasons may impose different selection pressures on organisms. Here, the authors propose that species may either maximize gains during the growth season or minimize losses during winter, and provide empirical support of such seasonal specialisation in two closely related butterfly species.

Whether net primary productivity in an aquatic ecosystem is limited by nitrogen (N), limited by phosphorus (P), or co-limited by N & P is determined by the relative supply of N and P to phytoplankton compared to their elemental requirements for primary production, often characterized by the “Redfield” ratio. The supply of these essential nutrients is affected by both external inputs and biogeochemical processes within the ecosystem. In this paper, we examine external sources of nutrients to aquatic systems and how the balance of N to P inputs influences nutrient limitation. For ocean subtropical gyres, a relatively balanced input of N and P relative to the Redfield ratio from deep ocean sources often leads to near co-limitation by N and P. For lakes, the external nutrient inputs come largely from watershed sources, and we demonstrate that on average the N:P ratio for these inputs across the United States is well above that needed by phytoplankton, which may contribute to P limitation in those lake that experience this average nutrient loading. Watershed inputs are also important for estuaries and coastal marine ecosystems, but ocean sources of nutrients are also significant contributors to overall nutrient loads. The ocean-nutrient sources of N and P are very often at or below the Redfield ratio of 16:1 molar, and can be substantially so, particularly in areas where the continental shelf is wide. This large input of coastal ocean nutrients with a low N:P ratio is one factor that may make N limitation more likely in many coastal marine ecosystems than in lakes.

Ruminants have a unique utilization of phosphate (Pi) based on the so-called endogenous Pi recycling to guarantee adequate Pi supply for ruminal microbial growth and for buffering short-chain fatty acids. Large amounts of Pi enter the gastrointestinal tract by salivary secretion. The high saliva Pi concentrations are generated by active secretion of Pi from blood into primary saliva via basolateral sodium (Na+)-dependent Pi transporter type II. The following subsequent intestinal absorption of Pi is mainly carried out in the jejunum by the apical located secondary active Na+-dependent Pi transporters NaPi IIb (SLC34A2) and PiT1 (SLC20A1). A reduction in dietary Pi intake stimulates the intestinal Pi absorption by increasing the expression of NaPi IIb despite unchanged plasma 1,25-dihydroxyvitamin D3 concentrations, which modulate Pi homeostasis in monogastric species. Reabsorption of glomerular filtrated plasma Pi is mainly mediated by the Pi transporters NaPi IIa (SLC34A1) and NaPi IIc (SLC34A3) in proximal tubule apical cells. The expression of NaPi IIa and the corresponding renal Na+-dependent Pi capacity were modulated by high dietary phosphorus (P) intake in a parathyroid-dependent manner. In response to reduced dietary Pi intake, the expression of NaPi IIa was not adapted indicating that renal Pi reabsorption in ruminants runs at a high level allowing no further increase when P intake is diminished. In bones and in the mammary glands, Na+-dependent Pi transporters are able to contribute to maintaining Pi homeostasis. Overall, the regulation of Pi transporter activity and expression by hormonal modulators confirms substantial differences between ruminant and non-ruminant species.

There has recently been significant interest in the concept of directly targeting intestinal phosphate transport to control hyperphosphatemia in patients with chronic kidney disease. However, we do not have a complete understanding of the cellular mechanisms that govern dietary phosphate absorption. Studies in the 1970s documented both active and passive pathways for intestinal phosphate absorption. However, following the cloning of the intestinal SLC34 cotransporter, NaPi-IIb, much of the research focused on the role of this protein in active transcellular phosphate absorption and the factors involved in its regulation. Generation of a conditional NaPi-IIb knockout mouse has demonstrated that this protein is critical for the maintenance of skeletal integrity during periods of phosphate restriction and that under normal physiological conditions, the passive sodium-independent pathway is likely be the more dominant pathway for intestinal phosphate absorption. The review aims to summarise the most recent developments in our understanding of the role of the intestine in phosphate homeostasis, including the acute and chronic renal adaptations that occur in response to dietary phosphate intake. Evidence regarding the overall contribution of the transcellular and paracellular pathways for phosphate absorption will be discussed, together with the clinical benefit of inhibiting these pathways for the treatment of hyperphosphatemia in chronic kidney disease.

In order to assess the progress toward eutrophication management goals, it is important to understand trends in land-based nutrient use. Here we present net anthropogenic nitrogen and phosphorus inputs (NANI and NAPI, respectively) for 2000 and 2010 for the Baltic Sea watershed. Overall, across the entire Baltic, between the 5-year periods centered on 2000 and 2010, NANI and NAPI decreased modestly by –6 and –4%, respectively, but with substantial regional variation, including major increases in the Gulf of Riga drainage basin (+19 and +58%, respectively) and decreases in the Danish Straits drainage basin (–25 and –40% respectively). The changes were due primarily to changes in mineral fertilizer use. Mineral fertilizers dominated inputs, at 57% of both NANI and NAPI in 2000, increasing to 68 and 70%, respectively, by 2010. Net food and feed imports declined over that period, corresponding to increased crop production; either fewer imports of food and feedstocks were required to feed humans and livestock, or more of these commodities were exported. A strong linear relationship exists between regional net nutrient inputs and riverine nutrient fluxes for both periods. About 17% of NANI and 4.7% of NAPI were exported to the sea in 2000; these relationships did not significantly differ from those for 2010. Changes in NANI from 2000 to 2010 across basins were directly proportional rather than linearly related to changes in total N (TN) fluxes to the sea (i.e., no change in NANI suggests no change in TN flux). Similarly, for all basins except those draining to the Baltic Proper, changes in NAPI were proportional to changes in total P (TP) fluxes. The Danish Straits decreased most between 2000 and 2010, where NANI and NAPI declined by 25 and 40%, respectively, and corresponding fluxes of TN and TP declined 31 and 18%, respectively. For the Baltic Proper, NAPI was relatively unchanged between 2000 and 2010, while riverine TP fluxes decreased 25%, due possibly to lagged effects of fertilizer reduction resulting from socio-political changes in the early 1990s or improvements in sewage treatment capabilities. For most regions, further reductions in NANI and NAPI could be achieved by more efficient production and greater substitution of manure for imported mineral fertilizers.

Polyphenism, the expression of discrete alternative phenotypes, is often a consequence of a developmental switch. physiological changes induced by a developmental switch potentially affect reaction norms, but the evolution and existence of alternative reaction norms remains poorly understood. Here, we demonstrate that, in the butterfly Pieris napi (Lepidoptera: Pieridae), thermal reaction norms of several life history traits vary adaptively among switch-induced alternative developmental pathways of diapause and direct development. The switch was affected both by photoperiod and temperature, ambient temperature during late development having the potential to override earlier photoperiodic cues. Directly developing larvae had higher development and growth rates than diapausing ones across the studied thermal gradient. Reaction norm shapes also differed between the alternative developmental pathways, indicating pathway-specific selection on thermal sensitivity. Relative mass increments decreased linearly with increasing temperature and were higher under direct development than diapause. Contrary to predictions, population phenology did not explain trait variation or thermal sensitivity, but our experimental design probably lacks power for finding subtle phenology effects. We demonstrate adaptive differentiation in thermal reaction norms among alternative phenotypes, and suggest that the consequences of an environmentally dependent developmental switch primarily drive the evolution of alternative thermal reaction norms in P. napi.

Phenotypic plasticity is produced and maintained by processes regulating the transcriptome. While differential gene expression is among the most important of these processes, relatively little is known about other sources of transcriptional variation. Previous work suggests that alternative splicing plays an extensive and functionally unique role in transcriptional plasticity, though plastically spliced genes may be more constrained than the remainder of expressed genes. In this study, we explore the relationship between expression and splicing plasticity, along with the genetic diversity in those genes, in an ecologically consequential polyphenism: facultative diapause. Using 96 samples spread over two tissues and 10 timepoints, we compare the extent of differential splicing and expression between diapausing and direct developing pupae of the butterfly Pieris napi. Splicing differs strongly between diapausing and direct developing trajectories but alters a smaller and functionally unique set of genes compared to differential expression. We further test the hypothesis that among these expressed loci, plastically spliced genes are likely to experience the strongest purifying selection to maintain seasonally plastic phenotypes. Genes with unique transcriptional changes through diapause consistently had the lowest nucleotide diversity, and this effect was consistently stronger among genes that were differentially spliced compared to those with just differential expression through diapause. Further, the strength of negative selection was higher in the population expressing diapause every generation. Our results suggest that maintenance of the molecular mechanisms involved in diapause progression, including post-transcriptional modifications, are highly conserved and likely to experience genetic constraints, especially in northern populations of P. napi.