Polar lipid composition of chloroplast thylakoids isolated from leaves grown under different lighting conditions

The polar acyl lipid composition was determined for samples of chloroplast thylakoids isolated from Pisum sativum plants grown at light intensities of 50 and 300 E·m^-2·s^-1 and from Aesculus hippocastanum leaves taken from shade or sun environments. Lighting conditions had no major effect on lipid class composition except for a small increase in the amount of monogalactosyldiacylglycerol relative to other lipids in low compared with high light and shade compared with sun conditions. The thylakoids from low light and shade environments also had, relative to those from high light and sun conditions, a substantial decrease in the level of trans-hexadecenoic acid in phosphatidyglycerol. In parallel with this there were lower lipid to chlorophyll ratios, higher overall fatty acid unsaturation, lower chlorophyll a to b ratios and increased relative levels of light harvesting chlorophyll a/b polypeptides as expected for an increase in the degree of thylakoid appression. With this in mind, our results on lipid class composition and content of trans-hexadecenoic acid are discussed in the context of the lateral distribution of lipids within the plane of membrane.Abbreviations DGDG digalactosyldiacylglycerol - EDTA ethylenediaminetetraacetic acid - HEPES N-2-hydroxyethylpiperazine-N-2-ethanesulfonic acid - LHC light harvesting chlorophyll a/b - MGDG monogalactosyldiacylglycerol - MPL minor phospholipids - PS1 photosystem one - PS2 photosystem two - SDS sodium dodecyl sulphate - SL sulphoquinovosyldiacylglycerol     

红树植物白骨壤叶片的解剖结构及其生态适应性

运用石蜡切片的方法对红树植物白骨壤的叶片进行解剖学的观察和研究。结果表明:白骨壤的叶为异面叶;上表皮为复表皮,有厚的角质膜及下皮层;下表皮上有大量的泌盐腺毛;栅栏组织多层,约占叶肉组织的2/3,海绵组织退化;机械组织和输导组织均很发达。反映了白骨壤叶片的结构与其生活的海生环境高度相适应。     

Iridoid glucosides and flavones from the aerial parts of Avicennia marina

Two new iridoid glucosides, namely, 2'-O-[(2E,4E)-5-phenylpenta-2,4-dienoyl]mussaenosidic acid (1; mussaenosidic acid = [1S-(1alpha,4aalpha,7alpha,7aalpha)]-1-(beta-D-glucopyranosyloxy)-1,4a,5,6,7,7a-hexahydro-7-hydroxy-7-methylcyclopenta[c]pyran-4-carboxylic acid) and 2'-O-(4-methoxycinnamoyl)mussaenosidic acid (2), were isolated from the aerial parts of the mangrove plant Avicennia marina. Beside that, one known iridoid glucoside, 2'-O-coumaroylmussaenosidic acid (3) and four known flavones (flavone = 2-phenyl-4H-1-benzopyran-4-one) including 4',5-dihydroxy-3',7-dimethoxyflavone (4), 4',5-dihydroxy-3',5',7-trimethoxyflavone (5), 4',5,7-trihydroxyflavone (6), and 3',4',5-trihydroxy-7-methoxyflavone (7) were also isolated and identified. The structures of these compounds were elucidated by NMR spectroscopy and by low- and high-resolution mass spectrometry. The chemotaxonomic significance of these findings was discussed. In addition, each isolated compound was evaluated for the ability of alpha,alpha-diphenyl-beta-picrylhydrazyl (DPPH) radical-scavenging activity.     

Gas exchange responses of a mangrove species, Avicennia marina, to waterlogged and drained conditions

This study was undertaken in summer on fully expanded leaves of Avicennia marina trees in the Beachwood Mangroves Nature Reserve, Durban,South Africa. Data sets were obtained over 5–7 days of relatively dry conditions and over two periods of 5 days during which the swamp was continuously inundated with dilute seawater (< 150 mol m⁻³NaCl). Gas exchange responses were strongly influenced by photosynthetic photon flux density (PPFD), leaf temperature and leaf to air vapour pressure deficit (Δw). Carbon dioxide exchange was saturated at a PPFD of about 800 μmol m⁻² s⁻¹. Maximal CO₂ exchange rates ranged from 8.5 to 9.9 μmol m⁻² s⁻¹ with no differences between drained and waterlogged conditions. Under drained conditions, leaf conductance,transpiration and internal CO₂ concentrations were generally lower, and water use efficiencies higher, than during waterlogging. Continuous waterlogging for 5 days had no adverse effect on CO₂ exchange. Xylem water potentials ranged from −1.32to −3.53 MPa during drained and from −1.02 to −2.65 Mpa during waterlogged conditions. These results are discussed in relation to anatomical and metabolic adaptations of A. marina to waterlogging stress. This revised version was published online in August 2006 with corrections to the Cover Date.     

镉胁迫对白骨壤幼苗钾钠镁吸收与分配的影响

采用砂基栽培,研究重金属镉(Cd)不同胁迫浓度(0、0.5、5、25、50、100和150mg·L-1)和胁迫时间(45和90 d)对红树植物白骨壤幼苗钾钠镁吸收与分配的影响.结果表明:白骨壤各器官对Cd的吸收和累积均表现为随Cd胁迫浓度上升而增加,随胁迫时间增加而增加的趋势;进入植物体内的Cd主要富集在根部,其次是凋落子叶,在150 mg·L-1浓度下分别占累积总量的66.9％和16.3％,这对减少Cd对植物全株的危害具有积极意义.受Cd胁迫影响90 d后,根和茎中Na含量呈上升趋势,叶和子叶中呈下降趋势;根和子叶中K含量呈下降趋势,茎和叶中变化不明显;Cd胁迫90 d后各器官(根、茎、叶和子叶)中Mg含量均低于对照,且与Cd含量呈显著负相关.     

Aerenchyma tissue development and gas-pathway structure in root of Avicennia marina (Forsk.) Vierh.

The aerenchyma differentiation in cable roots, pneumatophores, anchor roots, and feeding roots of the mangrove plant, Avicennia marina (Verbenaceae) was analyzed using a light microscope and scanning electron microscope. In all types, cortex cells were arranged in longitudinal columns extending from the endodermis to the epidermis. No cells in the cortex had intercellular spaces at the root tip (0–150 m), and aerenchyma started developing at 200 m from the root apex. The aerenchyma formation was due to cell separation (schizogeny) rather than cell lysis. The cell separation occurred between the longitudinal cell columns, forming long intercellular spaces along the root axis. During aerenchyma formation, the cortex cells enlarged longitudinally by 1.8–3.9 times and widened horizontally by 2.2–2.9 times. As a result, the aerenchyma had a pronounced tubular structure that was radially long, elliptical or oval in cross section and that ran parallel to the root axis. The tube had tapering ends, as did vessel elements, although there were no perforated plates. The interconnection between neighboring tubes was made by abundant small pores or canals that were schizogenous intercellular spaces between the wall cells. All aerenchyma tubes in the root were interconnected by these small pores serving as a gas pathway.     

Changes in the structure and the functional state of thylakoids under the conditions of osmotic shock

The effect of osmotic shock on the ultrastructure and functions of C-class pea chloroplasts has been examined. When incubated in a non-sucrose medium for 30 s or more, thylakoids were found to pass to a stable deformed state. This state was characterized by an altered orientation of thylakoids to each other with the lumen thickness remaining the same as in the normal state. Experiments with shorter incubation periods (10–20 s) revealed a swelling of thylakoids, which probably represented an intermediate stage. The deformation of the thylakoid system was accompanied by a decrease in the non-cyclic ATP synthesis but by an increase in the rate of cyclic photophosphorylation. Besides, the deformed thylakoids demonstrated an acceleration of the basal electron transport, as well a rise in the light-induced H⁺ and imidazol uptake. The data obtained are discussed in the light of membrane interactions fixing the configuration of a thylakoid.     

Relationship between chloroplast structure and O2 evolution rate of leaf discs in plants from different biotopes in South Finland

Abstract The chloroplast ultrastructure, especially the thylakoid organization, the polypeptide composition of the thylakoid membranes and photosynthetic O₂ evolution rate, chlorophyll (Chl) content and Chi a/b ratio were studied in leaves of nine plants growing in contrasting biotopes in the wild in South Finland. All the measurements were made at the beginning of the period of main growth on leaves approaching full expansion, when the CO₂-saturated O₂ evolution rate (measured at 20°C and 1500 μmol photons m^?2s^?1) was at a maximum, ranging from 19.2 to 6.9 μmol O₂ cm^?2 h^?1. Among the species, the Chi a/b ratio varied between 3.75 and 2.71. In the mesophyll chloroplasts, the ratio of the total length of appressed to non-appressed thylakoid membranes varied between 1.07 and 1.79, the number of partitions per granum varied between 2.8 and 12.0 and the grana area between 21 and 42% of the chloroplast area. There was a significant relationship between the rate of O₂ evolution of the leaf discs and the thylakoid organization in the mesophyll chloroplasts. The higher the O₂ evolution rate, the lower was the ratio of the total length of appressed to non-appressed thylakoid membranes and also the lower the grana area. Although the relationship of the photosynthetic rate with the Chi content and the Chi a/b ratio of the leaves was not as clear, a significant negative correlation existed between the Chi a/b ratio and the ratio of appressed to non-appressed thylakoid membranes, indicating lateral heterogeneity in the distribution of different Chl- protein complexes.     

Determination of in vivo oxygen uptake and carbon dioxide evolution rates from off-gas measurements under highly dynamic conditions

In vivo kinetics of Saccharomyces cerevisiae are studied, in a time window of 150 s, by analyzing the response of O(2) and CO(2) in the fermentor off-gas after perturbation of chemostat cultures by metabolite pulses. Here, a new mathematical method is presented for the estimation of the in vivo oxygen uptake rate (OUR) and carbon dioxide evolution rate (CER) directly from the off-gas data in such perturbation experiments. The mathematical construction allows effective elimination of delay and distortion in the off-gas measurement signal under highly dynamic conditions. A black box model for the fermentor off-gas system is first obtained by system identification, followed by the construction of an optimal linear filter, based on the identified off-gas model. The method is applied to glucose and ethanol pulses performed on chemostat cultures of S. cerevisiae. The estimated OUR is shown to be consistent with the independent dissolved oxygen measurement. The estimated in vivo OUR and CER provide valuable insights into the complex dynamic behavior of yeast and are essential for the establishment and validation of in vivo kinetic models of primary metabolism.     

Growth responses of the mangrove Avicennia marina to salinity: development and function of shoot hydraulic systems require saline conditions

Background and Aims Halophytic eudicots are characterized by enhanced growth under saline conditions. This study combines physiological and anatomical analyses to identify processes underlying growth responses of the mangrove Avicennia marina to salinities ranging from fresh- to seawater conditions.Methods Following pre-exhaustion of cotyledonary reserves under optimal conditions (i.e. 50 % seawater), seedlings of A. marina were grown hydroponically in dilutions of seawater amended with nutrients. Whole-plant growth characteristics were analysed in relation to dry mass accumulation and its allocation to different plant parts. Gas exchange characteristics and stable carbon isotopic composition of leaves were measured to evaluate water use in relation to carbon gain. Stem and leaf hydraulic anatomy were measured in relation to plant water use and growth.Key Results Avicennia marina seedlings failed to grow in 0–5 % seawater, whereas maximal growth occurred in 50–75 % seawater. Relative growth rates were affected by changes in leaf area ratio (LAR) and net assimilation rate (NAR) along the salinity gradient, with NAR generally being more important. Gas exchange characteristics followed the same trends as plant growth, with assimilation rates and stomatal conductance being greatest in leaves grown in 50–75 % seawater. However, water use efficiency was maintained nearly constant across all salinities, consistent with carbon isotopic signatures. Anatomical studies revealed variation in rates of development and composition of hydraulic tissues that were consistent with salinity-dependent patterns in water use and growth, including a structural explanation for low stomatal conductance and growth under low salinity.Conclusions The results identified stem and leaf transport systems as central to understanding the integrated growth responses to variation in salinity from fresh- to seawater conditions. Avicennia marina was revealed as an obligate halophyte, requiring saline conditions for development of the transport systems needed to sustain water use and carbon gain.     

Proteomic analysis of plasma membrane and tonoplast from the leaves of mangrove plant Avicennia officinalis

In order to understand the salt tolerance and secretion in mangrove plant species, gel electrophoresis coupled with LC‐MS‐based proteomics was used to identify key transport proteins in the plasma membrane (PM) and tonoplast fractions of Avicennia officinalis leaves. PM and tonoplast proteins were purified using two‐aqueous‐phase partitioning and density gradient centrifugation, respectively. Forty of the 254 PM proteins and 31 of the 165 tonoplast proteins identified were predicted to have transmembrane domains. About 95% of the identified proteins could be classified based on their functions. The major classes of proteins were predicted to be involved in transport, metabolic processes, defense/stress response, and signal transduction, while a few of the proteins were predicted to be involved in other functions such as membrane trafficking. The main classes of transporter proteins identified included H⁺‐ATPases, ATP‐binding cassette transporters, and aquaporins, all of which could play a role in salt secretion. These data will serve as the baseline membrane proteomic dataset for Avicennia species. Further, this information can contribute to future studies on understanding the mechanism of salt tolerance in halophytes in addition to salt secretion in mangroves. All MS data have been deposited in the ProteomeXchange with identifier PXD000837 ( http://proteomecentral.proteomexchange.org/dataset/PXD000837 ).     

Diversity and activity of aquatic fungi under low oxygen conditions

1. The objective was to test whether a decrease in oxygen concentration in streams affects the diversity and activity of aquatic hyphomycetes and consequently leaf litter decomposition. 2. Senescent leaves of Alnus glutinosa were immersed for 7 days in a reference stream, for fungal colonization, and then incubated for 18 days in microcosms at five oxygen concentrations (4%, 26%, 54%, 76% and 94% saturation). Leaf decomposition (as loss of leaf toughness), fungal diversity, reproduction (as spore production) and biomass (ergosterol content) were determined. 3. Leaf toughness decreased by 70% in leaves exposed to the highest O₂ concentration, whereas the decrease was substantially less (from 25% to 45%) in treatments with lower O₂. Fungal biomass decreased from 99 to 12 mg fungi g⁻¹ ash‐free dry mass on exposure to 94% and 4% O₂ respectively. Sporulation was strongly inhibited by reduction of dissolved O₂ in water (3.1 × 10⁴ versus 1.3 × 10³ spores per microcosms) for 94% and 4% saturation respectively. 4. A total of 20 species of aquatic hyphomycetes were identified on leaves exposed to 94% O₂, whereas only 12 species were found in the treatment with 4% O₂ saturation. Multidimensional scaling revealed that fungal assemblages exposed to 4% O₂ were separated from all the others. Articulospora tetracladia, Cylindrocarpon sp. and Flagellospora curta were the dominant species in microcosms with 4% O₂, while Flagellospora curvula and Anguillospora filiformis were dominant at higher O₂ concentrations. 5. Overall results suggest that the functional role of aquatic hyphomycetes as decomposers of leaf litter is limited when the concentration of dissolved oxygen in streams is low.     

Fluoride inhibition of oxygen evolution; new evidence from Cl-NMR measurements

³⁵Cl-NMR was used to investigate the nature of F⁻-induced inhibition of oxygen-evolution in thylakoids from the mangrove, Avicennia marina. These studies showed a correlation between F⁻ inhibition of oxygen evolving activity and increased bulk Cl⁻ relaxation, possibly associated with the formation of a new class of high-affinity Cl⁻-binding sites, or a change in the nature of the existing binding sites, induced by F⁻. The presence of added Cl⁻ did not alter the F⁻-induced inhibition of oxygen evolution. Increased Cl⁻ relaxation and F⁻-induced inhibition of oxygen evolution occurred at lower concentrations of F⁻ at pH 7.8 than at pH 6.3. In mangrove thylakoids. F⁻-induced inhibition of oxygen evolution does not appear to be due to competition with Cl⁻ for Cl⁻ binding sites, but instead involves some other interaction close to the oxygen-evolving complex.     

Relations between electron transport rates determined by pulse amplitude modulated chlorophyll fluorescence and oxygen evolution in macroalgae under different light conditions

The relationship between O₂-based gross photosynthesis (GP) and in vivo chlorophyll fluorescence of Photosystem II-based electron transport rate (ETR) as well as the relationship between effective quantum yield of fluorescence (Φ_PSII) and quantum yield of oxygen evolution (Φ_{O_2}) were examined in the green algae Ulva rotundata and Ulva olivascens and the red alga Porphyra leucosticta collected from the field and incubated for 3 days at 100 μmol m⁻² s⁻¹ in nutrient enriched seawater. Maximal GP was twice as high in Ulva species than that measured in P. leucosticta. In all species ETR was saturated at much higher irradiance than GP. The initial slope of ETR versus absorbed irradiance was higher than that of GP versus absorbed irradiance. Only under absorbed irradiances below saturation or at values of GP <2 μmol O₂ m⁻² s⁻¹ a linear relationship was observed. In the linear phase, calculated O₂ evolved /ETR molar ratios were closed to the theoretical value of 0.25 in Ulva species. In P. leucosticta, the estimated GP was associated to the estimated ETR only at high irradiances. ETR was determined under white light, red light emitting by diodes and solar radiation. In Ulva species the maximal ETR was reached under red light and solar radiation whereas in P. leucosticta the maximal ETR was reached under white light and minimal under red light. These results are in agreement with the known action spectra for photosynthesis in these species. In the case of P. leucosticta, GP and ETR were additionally determined under saturating irradiance in algae pre-incubated for one week under white light at different irradiances and at white light (100 μmol m⁻² s⁻¹) enriched with far-red light. GP and growth rate increased at a growth irradiance of 500 μmol m⁻² s⁻¹ becoming photoinhibited at higher irradiances, while ETR increased when algae were exposed to the highest growth irradiance applied (2000 μmol m⁻² s⁻¹). The calculated O₂ evolved /ETR molar ratios were close to the theoretical value of 0.25 when algae were pre-incubated under 500–1000 μmol m⁻² s⁻¹. The enrichment by FR light provoked a decrease in both GP and ETR and an increase of nonphotochemical quenching although the irradiance of PAR was maintained at a constant level. In addition to C assimilation, other electron sinks, such as nitrogen assimilation, affected the GP–ETR relationship. The slopes of GP versus ETR or Φ_PSII versus Φ_{O_2} were lower in the algae with the highest N assimilation capacity, estimated as nitrate reductase activity and internal nitrogen contents, i.e., Ulva rotundata and Porphyra leucosticta, than that observed in U. olivascens. The possible mechanisms to explain this discrepancy between GP and ETR are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Changes in lipid composition of Photosystem 1 particles from thylakoids phosphorylated under reductive or anaerobic conditions

Changes in lipid composition of Photosystem 1 (PS 1) particles isolated from thylakoids phosphorylated under reductive or anaerobic conditions have been studied. Under reductive conditions, there was an increase in monogalactosyldiacylglycerol containing highly saturated fatty acids and phosphatidylglycerol containing transhexadecenoic fatty acid. Under anaerobic conditions, the amount of all lipid classes was increased. As we have shown earlier (S. V. Manuilskaya, O. I. Volovik, A. I. Mikhno, A. I. Polischuk and S. M. Kochubey (1990) Photosynthetica 24: 419–423) these changes were due to a co-migration of some lipid species and light-harvesting chlorophyll a/b complex LHC II from PS 2 to PS 1. These data allow us to conclude that LHC II consists of the lipoproteins containing specific lipids. Different composition of lipids co-migrating with LHC II under various conditions of phosphorylation might be caused by the variety of LHC II subpopulations transferred under each reductive condition.Abbreviations PS 1 Photosystem 1 - PS 2 Photosystem 2 - LHC II light-harvesting chlorophyll a/b protein complex II - Chl chlorophyll - MGDG monogalactosyldiacylglycerol - DGDG digalactosyldiacylglycerol - PG phosphatidylglycerol - SQDG sulfoquinovosyldiacylglycerol     

High misses after odd flashes in oxygen evolution in thoroughly dark-adapted thylakoids from pea and Chenopodium album

In Photosystem II (PS II), water is oxidized to molecular oxygen and plastoquinone is reduced to plastoquinol. The oxidation of water requires the accumulation of four oxidizing equivalents, through the so-called S-states of the oxygen evolving complex; the production of plastoquinol requires the accumulation of two reducing equivalents on a bound plastoquinone, Q_B. It has been generally believed that during the flash-induced transition of each of the S-states (S_n S_n+1, where n=0, 1, 2 and 3), a certain small but equal fraction of the PS II reaction centers are unable to function and, thus, miss being turned over. We used thoroughly dark-adapted thylakoids from peas (Pisum sativum) and Chenopodium album (susceptible and resistant to atrazine) starting with 100% of the oxygen evolving complex in the S₁ state. Thylakoids were illuminated with saturating flashes, providing a double hit parameter of about 0.07. Our experimental data on flashnumber dependent oscillations in the amount of oxygen per flash fit very well with a binary pattern of misses: 0, 0.2, 0, 0.4 during S₀ S₁, S₁ S₂, S₂ S₃ and S₃ S₀ transitions. Addition of 2 mM ferricyanide appears to shift this pattern by one flash. These results are consistent with the bicycle model recently proposed by V. P. Shinkarev and C. A. Wraight (Oxygen evolution in photosynthesis: From unicycle to bicycle, 1993, Proc Natl Acad Sci USA 90: 1834–1838), where misses are due to the presence of P⁺ or Q_A ^- among the various equilibrium states of PS II centers.Abbreviations miss parameter - double hit parameter - PS II Photosystem II - Q_A primary one-electron acceptor of PS II, a plastoquinone molecule - Q_B secondary plastoquinone two-electron acceptor of PS II - S-states (S_n, where n=0, 1, 2, 3 or 4) redox states of the oxygen evolving complex     

钝顶螺旋藻在不同光照条件下的放氧特性

钝顶螺旋藻在持续照光和中等频率 (0.01~20 Hz) 的光/暗交替照光下的放氧特性对光生物反应器的设计和操作具有重要意义。构建了一套可实现光/暗交替的光生物反应器系统对此进行研究,结果显示：根据与放氧速率的关系,可以将光强分为4个区：光限制区 (0~335 μmol/(m2·s)),过渡区 (335~875 μmol/(m2·s)),光饱和区 (875~2 775 μmol/(m2·s)) 以及光抑制区 (2 775 μmol/(m2·s)以上)。提高光/暗频率能否提高微藻光合速率取决于所采用的光强和     

Modelling reveals endogenous osmotic adaptation of storage tissue water potential as an important driver determining different stem diameter variation patterns in the mangrove species Avicennia marina and Rhizophora stylosa

Background

Stem diameter variations are mainly determined by the radial water transport between xylem and storage tissues. This radial transport results from the water potential difference between these tissues, which is influenced by both hydraulic and carbon related processes. Measurements have shown that when subjected to the same environmental conditions, the co-occurring mangrove species Avicennia marina and Rhizophora stylosa unexpectedly show a totally different pattern in daily stem diameter variation.

Methods

Using in situ measurements of stem diameter variation, stem water potential and sap flow, a mechanistic flow and storage model based on the cohesion–tension theory was applied to assess the differences in osmotic storage water potential between Avicennia marina and Rhizophora stylosa.

Key results

Both species, subjected to the same environmental conditions, showed a resembling daily pattern in simulated osmotic storage water potential. However, the osmotic storage water potential of R. stylosa started to decrease slightly after that of A. marina in the morning and increased again slightly later in the evening. This small shift in osmotic storage water potential likely underlaid the marked differences in daily stem diameter variation pattern between the two species.

Conclusions

The results show that in addition to environmental dynamics, endogenous changes in the osmotic storage water potential must be taken into account in order to accurately predict stem diameter variations, and hence growth.     

Atmospheric oxygen level and the evolution of insect body size

Insects are small relative to vertebrates, possibly owing to limitations or costs associated with their blind-ended tracheal respiratory system. The giant insects of the late Palaeozoic occurred when atmospheric PO₂ (aPO₂) was hyperoxic, supporting a role for oxygen in the evolution of insect body size. The paucity of the insect fossil record and the complex interactions between atmospheric oxygen level, organisms and their communities makes it impossible to definitively accept or reject the historical oxygen-size link, and multiple alternative hypotheses exist. However, a variety of recent empirical findings support a link between oxygen and insect size, including: (i) most insects develop smaller body sizes in hypoxia, and some develop and evolve larger sizes in hyperoxia; (ii) insects developmentally and evolutionarily reduce their proportional investment in the tracheal system when living in higher aPO₂, suggesting that there are significant costs associated with tracheal system structure and function; and (iii) larger insects invest more of their body in the tracheal system, potentially leading to greater effects of aPO₂ on larger insects. Together, these provide a wealth of plausible mechanisms by which tracheal oxygen delivery may be centrally involved in setting the relatively small size of insects and for hyperoxia-enabled Palaeozoic gigantism.     

On the co‐evolution of surface oxygen levels and animals

Few topics in geobiology have been as extensively debated as the role of Earth's oxygenation in controlling when and why animals emerged and diversified. All currently described animals require oxygen for at least a portion of their life cycle. Therefore, the transition to an oxygenated planet was a prerequisite for the emergence of animals. Yet, our understanding of Earth's oxygenation and the environmental requirements of animal habitability and ecological success is currently limited; estimates for the timing of the appearance of environments sufficiently oxygenated to support ecologically stable populations of animals span a wide range, from billions of years to only a few million years before animals appear in the fossil record. In this light, the extent to which oxygen played an important role in controlling when animals appeared remains a topic of debate. When animals originated and when they diversified are separate questions, meaning either one or both of these phenomena could have been decoupled from oxygenation. Here, we present views from across this interpretive spectrum—in a point–counterpoint format—regarding crucial aspects of the potential links between animals and surface oxygen levels. We highlight areas where the standard discourse on this topic requires a change of course and note that several traditional arguments in this “life versus environment” debate are poorly founded. We also identify a clear need for basic research across a range of fields to disentangle the relationships between oxygen availability and emergence and diversification of animal life.