RESUMO
[This corrects the article DOI: 10.3389/fphys.2017.00702.].
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Survival under stress, such as exposure to hypoxia, anoxia, freezing, dehydration, air exposure of water breathing organisms, and estivation, is commonly associated to enhanced endogenous antioxidants, a phenomenon coined "preparation for oxidative stress" (POS). The regulation of free radical metabolism seems to be crucial under these selective pressures, since this response is widespread among animals. A hypothesis of how POS works at the molecular level was recently proposed and relies on two main processes: increased reactive species production under hypoxia, and activation of redox-sensitive transcription factors and signaling pathways, increasing the expression of antioxidants. The present paper brings together the current knowledge on POS and considers its future directions. Data indicate the presence of POS in 83 animal species (71.6% among investigated species), distributed in eight animal phyla. Three main research challenges on POS are presented: (i) to identify the molecular mechanism(s) that mediate/induce POS, (ii) to identify the evolutionary origins of POS in animals, and (iii) to determine the presence of POS in natural environments. We firstly discuss the need of evidence for increased RS production in hypoxic conditions that underlie the POS response. Secondly, we discuss the phylogenetic origins of POS back 700 million years, by identifying POS-positive responses in cnidarians. Finally, we present the first reports of the POS adaptation strategy in the wild. The investigation of these research trends and challenges may prove useful to understand the evolution of animal redox adaptations and how they adapt to increasing stressful environments on Earth.
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A key question in evolutionary genetics is why certain mutations or certain types of mutation make disproportionate contributions to adaptive phenotypic evolution. In principle, the preferential fixation of particular mutations could stem directly from variation in the underlying rate of mutation to function-altering alleles. However, the influence of mutation bias on the genetic architecture of phenotypic evolution is difficult to evaluate because data on rates of mutation to function-altering alleles are seldom available. Here, we report the discovery that a single point mutation at a highly mutable site in the ß(A)-globin gene has contributed to an evolutionary change in hemoglobin (Hb) function in high-altitude Andean house wrens (Troglodytes aedon). Results of experiments on native Hb variants and engineered, recombinant Hb mutants demonstrate that a nonsynonymous mutation at a CpG dinucleotide in the ß(A)-globin gene is responsible for an evolved difference in Hb-O2 affinity between high- and low-altitude house wren populations. Moreover, patterns of genomic differentiation between high- and low-altitude populations suggest that altitudinal differentiation in allele frequencies at the causal amino acid polymorphism reflects a history of spatially varying selection. The experimental results highlight the influence of mutation rate on the genetic basis of phenotypic evolution by demonstrating that a large-effect allele at a highly mutable CpG site has promoted physiological differentiation in blood O2 transport capacity between house wren populations that are native to different elevations.
Assuntos
Adaptação Biológica/genética , Altitude , Hemoglobinas/metabolismo , Fenótipo , Mutação Puntual/genética , Aves Canoras/genética , Globinas beta/genética , Adaptação Biológica/fisiologia , Animais , Sequência de Bases , Clonagem Molecular , Hemoglobinas/genética , Hemoglobinas/isolamento & purificação , Dados de Sequência Molecular , Taxa de Mutação , Oxigênio/metabolismo , Peru , Reação em Cadeia da Polimerase , Proteínas Recombinantes/metabolismo , Análise de Sequência de DNA , Aves Canoras/fisiologia , Espectrometria de Massas em TandemRESUMO
The non-ionized form of ammonia is very toxic to many aquatic species. It is especially important in several aspects of fish biology. A large range of organismal strategies for coping with environmental stressors is usually observed in living organisms. Among those, the responses for managing chemical stressors are well studied. The present work compares biochemical responses of two evolutionarily close species, Hoplias malabaricus and Hoplerythrinus unitaeniatus, exposed to environmental ammonia. Adult fish were submitted to 1.0 mg/L of ammonium chloride for 24 hours, and plasma ammonia and urea levels were determined. The activities of OUC enzymes OCT and ARG, and the accessory enzyme GS, were quantified in liver extract and are expressed below in mumol/min/mg of wet tissue. Increases in OUC enzymes (GS from 1.14 to 2.43, OCT from 0.81 to 1.72, and ARG from 3.15 to 4.23), plasma ammonia (from 0.95 to 1.42 mmol/L), and plasma urea (from 0.82 to 1.53 mmol/L) were observed (p 0.05) in H. malabaricus exposed to 1 mg/L of ammonia chloride. The GS in H. unitaeniatus increased from 1.43 to 1.84, however the OCT, ARG, and plasma urea from H. unitaeniatus did not change. These data indicate that each species responds differently to the same environmental stressor.
A forma não ionizada da amônia é muito tóxica a vários organismos aquáticos, sendo particularmente importante em muitos aspectos da biologia dos peixes. Um amplo grupo de estratégias para enfrentar os estressores ambientais pode ser observado nos organismos vivos. Dentre estas, as respostas aos estressores químicos são bem estudadas. O presente trabalho compara respostas bioquímicas de duas espécies evolutivamente próximas Hoplias malabaricus e Hoplerythrinus unitaeniatus, expostas à amônia ambiental. Peixes adultos foram submetidos a 1.0 mg/L de cloreto de amônio por 24 horas e foram determinados os níveis plasmáticos de amônia e uréia. As atividades das enzimas do COU, OCT e ARG e a enzima acessória GS foram quantificadas em extrato de fígado e são expressas em mimol/min/mg de tecido úmido. Foi observado em H. malabaricus, exposto a 1,0 mg/L de cloreto de amônio, aumento (p 0,05) nas enzimas: GS, de 1,14 para 2,43; OCT, de 0,81 para 1,72; ARG, de 3,15 para 4,23; na amônia plasmática, de 0,95 para 1,42 mmol/L, e na uréia plasmática, de 0,82 para 1,53. A GS de H. malabaricus aumentou de 1,43 para 1,84, todavia, OCT, ARG e uréia plasmática não variaram. Esses dados mostram que ambas as espécies, taxonomicamente próximas, respondem distintamente ao mesmo estressor ambiental.
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Oxygen of tropical freshwater environments fluctuates drastically. Eutrophic lakes and ponds of warm waters frequently reach very low oxygen concentrations. This is the most common habitat of the banded knife fish "tuvira" Gymnotus carapo. This electric fish is reported to present bimodal breathing to cope with low environmental oxygen. Biochemical responses can be also observed in fishes facing hypoxia but none were studied in tuvira. In the present study, haematological and metabolic changes were investigated in two groups of fish exposed to hypoxia for 1 and 3 hours. Haematocrit, red blood cells and haemoglobin concentration indicated erythrocyte release from hematopoietic organs and swelling of red blood cells. Glycogen, glucose, lactate, pyruvate, and amino acids were quantified in liver, kidney and white muscle. The metabolic profile of G. carapo to cope with hypoxia suggested liver gluconeogenesis probably supported by proteolysis. The kidney and liver presented the same biochemical trend suggesting similar metabolic role for both organs. Glucogenolysis followed by glucose fermentation and protein mobilisation was observed in the white muscle. The air breathing behaviour of tuvira works in parallel with metabolism to prevent damages from hypoxia. Metabolic adjustments are observed when the air taking is avoided.
O oxigênio de água doce dos ambientes tropicais flutua drasticamente. Lagos e lagoas eutróficos de ambientes temperados freqüentemente atingem baixas concentrações de oxigênio. Este é o habitat mais comum da tuvira Gymnotus carapo. Neste peixe elétrico é descrita a respiração bimodal para enfrentar baixos níveis de oxigênio. Em peixes, as respostas bioquímicas também podem ser observadas, mas nenhuma foi estudada em tuvira. Neste estudo foram investigadas as alterações hematológicas e metabólicas em dois grupos de peixes expostos à hipóxia por 1 e 3 horas. O hematócrito, os eritrócitos e a concentração de hemoglobina indicaram liberação de hemácias por órgãos hematopoiéticos e intumescimento celular. Glicogênio, glicose, lactato, piruvato e aminoácidos foram quantificados no fígado, no rim e em músculo branco. O perfil metabólico de G. carapo para enfrentar a hipóxia sugeriu neoglicogênese hepática por proteólise. O rim e o fígado apresentaram a mesma tendência metabólica, sugerindo o mesmo papel metabólico de ambos os órgãos em hipóxia. Foi observada neoglicogênese em músculo branco, seguida da fermentação de glicose e da mobilização protéica. O comportamento de respiração área em tuvira funciona paralelamente ao metabolismo adaptativo para prevenir os danos da hipóxia. Os ajustes metabólicos foram observados a partir do momento em que a tomada de ar foi bloqueada.
RESUMO
Oxygen of tropical freshwater environments fluctuates drastically. Eutrophic lakes and ponds of warm waters frequently reach very low oxygen concentrations. This is the most common habitat of the banded knife fish "tuvira" Gymnotus carapo. This electric fish is reported to present bimodal breathing to cope with low environmental oxygen. Biochemical responses can be also observed in fishes facing hypoxia but none were studied in tuvira. In the present study, haematological and metabolic changes were investigated in two groups of fish exposed to hypoxia for 1 and 3 hours. Haematocrit, red blood cells and haemoglobin concentration indicated erythrocyte release from hematopoietic organs and swelling of red blood cells. Glycogen, glucose, lactate, pyruvate, and amino acids were quantified in liver, kidney and white muscle. The metabolic profile of G. carapo to cope with hypoxia suggested liver gluconeogenesis probably supported by proteolysis. The kidney and liver presented the same biochemical trend suggesting similar metabolic role for both organs. Glucogenolysis followed by glucose fermentation and protein mobilisation was observed in the white muscle. The air breathing behaviour of tuvira works in parallel with metabolism to prevent damages from hypoxia. Metabolic adjustments are observed when the air taking is avoided.
O oxigênio de água doce dos ambientes tropicais flutua drasticamente. Lagos e lagoas eutróficos de ambientes temperados freqüentemente atingem baixas concentrações de oxigênio. Este é o habitat mais comum da tuvira Gymnotus carapo. Neste peixe elétrico é descrita a respiração bimodal para enfrentar baixos níveis de oxigênio. Em peixes, as respostas bioquímicas também podem ser observadas, mas nenhuma foi estudada em tuvira. Neste estudo foram investigadas as alterações hematológicas e metabólicas em dois grupos de peixes expostos à hipóxia por 1 e 3 horas. O hematócrito, os eritrócitos e a concentração de hemoglobina indicaram liberação de hemácias por órgãos hematopoiéticos e intumescimento celular. Glicogênio, glicose, lactato, piruvato e aminoácidos foram quantificados no fígado, no rim e em músculo branco. O perfil metabólico de G. carapo para enfrentar a hipóxia sugeriu neoglicogênese hepática por proteólise. O rim e o fígado apresentaram a mesma tendência metabólica, sugerindo o mesmo papel metabólico de ambos os órgãos em hipóxia. Foi observada neoglicogênese em músculo branco, seguida da fermentação de glicose e da mobilização protéica. O comportamento de respiração área em tuvira funciona paralelamente ao metabolismo adaptativo para prevenir os danos da hipóxia. Os ajustes metabólicos foram observados a partir do momento em que a tomada de ar foi bloqueada.