Darwinian evolution does not rule out the gaia hypothesis

This study explores so-called Darwinian Daisyworlds mathematically rigorously in detail. The original Daisyworld was introduced by Watson & Lovelock (1983) to demonstrate how two species of daisies regulate the global temperature of their planet through competition among these species against the rising solar luminosity, i.e. the Gaia hypothesis. Its variants are Darwinian Daisyworlds in which daisies can adapt themselves to the local temperature. Robertson & Robinson (1998) insist their Darwinian daisies lose the ability for temperature regulation on the basis of their spreadsheet simulations. Lenton & Lovelock (2000) point out that the constraints on adaptation recovers Darwinian daisies' ability of temperature regulation on the basis of their Euler-code simulations. The present study shows there exist the exact and closed-form solutions to these two Daisyworlds. The results contradict the former studies: Robertson and Robinson's daisies do regulate the global temperature even longer than non-adaptive daisies; Lenton and Lovelock's daisies are less adaptive than Robertson and Robinson's daisies because of the constraints on adaptation; the introduction of weak adaptability drives species into a dead end of evolution. Thus, the present results confirm that the Gaia hypothesis and Darwinian evolution can coexist.     

Catastrophic desert formation in Daisyworld

Feedback between life and its environment is ubiquitous but the strength of coupling and its global implications remain hotly debated. Abrupt changes in the abundance of life for small changes in forcing provide one indicator of regulation, for example, when vegetation-climate feedback collapses in the formation of a desert. Here we use a two-dimensional "Daisyworld" model with curvature to show that catastrophic collapse of life under gradual forcing provides a testable indicator of environmental feedback. When solar luminosity increases to a critical value, a desert forms across a wide band of the planet. The scale of collapse depends on the strength of feedback. The efficiency of temperature regulation is limited by mutation rate in an analogous manner to the limitation of adaptive fitness in evolutionary theories. The final state of the system emerging from single-site rules can be described by two global quantities: optimization of temperature regulation and maximization of diversity, which are mathematically analogous to energy and entropy in thermodynamics.     

One-dimensional daisyworld: spatial interactions and pattern formation

The zero-dimensional daisyworld model of Watson and Lovelock (1983) demonstrates that life can unconsciously regulate a global environment. Here that model is extended to one dimension, incorporating a distribution of incoming solar radiation and diffusion of heat consistent with a spherical planet. Global regulatory properties of the original model are retained. The daisy populations are initially restricted to hospitable regions of the surface but exert both global and local feedback to increase this habitable area, eventually colonizing the whole surface. The introduction of heat diffusion destabilizes the coexistence equilibrium of the two daisy types. In response, a striped pattern consisting of blocks of all black or all white daisies emerges. There are two mechanisms behind this pattern formation. Both are connected to the stability of the system and an overview of the mathematics involved is presented. Numerical experiments show that this pattern is globally determined. Perturbations in one region have an impact over the whole surface but the regulatory properties of the system are not compromised by transient perturbations. The relevance of these results to the Earth and the wider climate modelling field is discussed.     

Symbiotic physiology promotes homeostasis in Daisyworld

A connection is hypothesized between the physiological consequences of mutualistic symbiosis and life's average long-term impact on certain highly biologically conserved environmental variables. This hypothesis is developed analytically and with a variant of the Daisyworld model. Biological homeostasis is frequently effective due to co-ordination between opposing physiological “rein” functions, which buffer an organism in response to an external (often environmental) perturbation. It is proposed that during evolutionary history the pooling of different species' physiological functions in mutualistic symbioses increased the range of suboptimal environmental conditions that could be buffered against—a mutual tolerance benefit sometimes sufficient to outweigh the cost of cooperation. A related argument is that for a small number of biologically-crucial physical variables (i) the difference between organism interiors and the life-environment interface is relatively low, and (ii) the biologically optimum level of that variable is relatively highly conserved across different species. For such variables, symbiosis tends to cause (at a cost) an increase in the number of environmental buffering functions per unit of selection, which in turn biases the overall impact of the biota on the state of the variable towards the biological optimum. When a costly but more temperature-tolerant and physiologically versatile symbiosis between one black (warming) and one white (cooling) “daisy” is added to the (otherwise unaltered) Daisyworld parable, four new results emerge: (1) The extension of habitability to a wider luminosity range, (2) resistance to the impact of “cheater” white daisies with cold optima, that derive short-term benefit from environmental destabilisation, (3) the capacity to maintain residual, oscillatory regulation in response to forcings that change more rapidly than allele frequencies and (crucially) (4) “succession”-type dynamics in which the tolerant symbiosis colonises and to an extent makes habitable an otherwise lifeless environment, but is later displaced by free-living genotypes that have higher local fitness once conditions improve. The final result is arguably analogous to lichen colonisation of the Neoproterozoic land surface, followed by the Phanerozoic rise of vascular plants. Caution is necessary in extrapolating from the Daisyworld parable to real ecology/geochemistry, but sufficiently conserved variables may be water potential, macronutrient stoichiometry and (to a lesser extent) the temperature window for metabolic activity.     

Topographic,latitudinal and climatic distribution of Pinus coulteri: geographic range limits are not at the edge of the climate envelope

With changing climate, many species are projected to move poleward or to higher elevations to track suitable climates. The prediction that species will move poleward assumes that geographically marginal populations are at the edge of the species' climatic range. We studied Pinus coulteri from the center to the northern (poleward) edge of its range, and examined three scenarios regarding the relationship between the geographic and climatic margins of a species' range. We used herbarium and iNaturalist.org records to identify P. coulteri sites, generated a species distribution model based on temperature, precipitation, climatic water deficit, and actual evapotranspiration, and projected suitability under future climate scenarios. In fourteen populations from the central to northern portions of the range, we conducted field studies and recorded elevation, slope and aspect (to estimate solar insolation) to examine relationships between local and regional distributions. We found that northern populations of P. coulteri do not occupy the cold or wet edge of the species' climatic range; mid‐latitude, high elevation populations occupy the cold margin. Aspect and insolation of P. coulteri populations changed significantly across latitudes and elevations. Unexpectedly, northern, low‐elevation stands occupy north‐facing aspects and receive low insolation, while central, high‐elevation stands grow on more south‐facing aspects that receive higher insolation. Modeled future climate suitability is projected to be highest in the central, high elevation portion of the species range, and in low‐lying coastal regions under some scenarios, with declining suitability in northern areas under most future scenarios. For P. coulteri, the lack of high elevation habitat combined with a major dispersal barrier may limit northward movement in response to a warming climate. Our analyses demonstrate the importance of distinguishing geographically vs. climatically marginal populations, and the importance of quantitative analysis of the realized climate space to understand species range limits.     

Daisyworld inhabited with daisies incorporating a seed size/number trade-off: the mechanism of negative feedback on selection from a standpoint of the competition theory

We reexamined a Daisyworld model from the traditional view of competition theory. Unlike the original model, white and black daisies in our model incorporate a seeding/germination trade-off against bare ground area without assuming the local temperature reward. As a result, the planetary temperature is automatically regulated by two species if the following conditions are met: (i) the species react equally to an environmental condition, but one can alter the environmental condition in the opposite direction to the other. (ii) that one of the two cannot have both a higher maximal growth rate (mu(max)) and lower half-saturation constant (K) than those of the other. In other words, a pair of phenotypes incorporates a trade-off between quality and number of seeds. We found that the homeostatic regulation can also be reconciled with the adaptive evolution of optimal temperature. The results of simulation imply that biotic environmental feedback can also be maintained when the emergence of polymorphisms (black and white daisies) is closely linked to such a trade-off.     

A fitness based analysis of Daisyworld

The Gaia hypothesis [Lovelock, J., Margulis, L., 1974. Atmospheric homeostasis: the Gaia hypothesis. Tellus 26, 1], that the earth functions as a self-regulating system, has never sat particularly comfortably with ideas in mainstream biology [Anon, 2002. In pursuit of arrogant simplicities. Nature 416, 247]. A lack of any clear role for evolution in the model has led to claims of teleology-that self-regulation emerges because it is pre-ordained to do so [Doolittle, W.F., 1981. Is nature really motherly? CoEvol. Q. 58-63; Dawkins, R., 1979. The Extended Phenotype. Oxford University Press, Oxford]. The Daisyworld parable [Watson, A.J., Lovelock, J.E., 1983. Biological homeostasis of the global environment--the parable of Daisyworld. Tellus B 35, 284], a simple mathematical illustration of Gaia, went some way to addressing these critiques but, despite recent success in incorporating natural selection [Stocker, S.,1995. Regarding mutations in Daisyworld models. J. Theor. Biol. 175, 495; Lenton, T.M., 1998. Gaia and natural selection. Nature 394, 439; Lenton, T.M., Lovelock, J.E., 2001. Daisyworld revisited: quantifying biological effects on planetary self-regulation. Tellus B 53, 288; Wood, A.J., Ackland, G.J., Lenton, T.M., 2006. Mutation of albedo and growth response leads to oscillations in a spatial Daisyworld. J. Theor. Biol. 242, 188], it remains a widely held view that the ideas are inconsistent with biological principles. We show that standard methodology from quantitative genetics can be used to predict the stationary states and dynamic behaviour of Daisyworlds. The system regulates its temperature due to the low-level evolutionary dynamics of competition between the thermally coupled daisies, no higher level principle is invoked. A reconciliation of Gaia with evolutionary theory may allow further development of evolutionary arguments for the existence of global self-regulatory systems.     

Temporal trends and variability in a high-arctic ecosystem in Greenland: multidimensional analyses of limnic and terrestrial ecosystems

The high arctic is undergoing a faster change in climate than most other regions of the planet, with already observed ecological consequences. Combined with the characteristics of high-arctic ecosystems, such as low species redundancy, high seasonality and weather extremes, shifts in individual species performance and phenology may lead to altered interaction dynamics through trophic mismatch and cascades. An ecosystem approach is therefore desirable in the attempt to understand the multidimensional impacts of climate. Here, we present ecosystem-wide trend analyses of a long-term dataset on terrestrial and limnic biota with focus on the distribution of observed trends and associated variation across the ecosystem. We used 114 time series drawn from 11 abiotic variables, 19 terrestrial and 7 limnic biotic species/taxa and compared temporal trends, changes and abrupt shifts in the variation within and across the two biota. A total of 36 % of the time series analysed showed a significant trend during the study period with a higher frequency of trends occurring within performance variables. Overall, the changes tended to be negative, indicating advances in phenology but reduced species performance. General system variance was also higher in the limnic biota than in the terrestrial biota, both exhibiting increasing variance up through the trophic system. Overall, our results suggest that multiple biotic responses to the climatic changes in this high-arctic ecosystem are not synchronised across trophic levels and may differ qualitatively and quantitatively between terrestrial and limnic biota.     

Ice-age survival of Atlantic cod: agreement between palaeoecology models and genetics

Scant scientific attention has been given to the abundance and distribution of marine biota in the face of the lower sea level, and steeper latitudinal gradient in climate, during the ice-age conditions that have dominated the past million years. Here we examine the glacial persistence of Atlantic cod (Gadus morhua) populations using two ecological-niche-models (ENM) and the first broad synthesis of multi-locus gene sequence data for this species. One ENM uses a maximum entropy approach (Maxent ); the other is a new ENM for Atlantic cod, using ecophysiological parameters based on observed reproductive events rather than adult distribution. Both the ENMs were tested for present-day conditions, then used to hindcast ranges at the last glacial maximum (LGM) ca 21kyr ago, employing climate model data. Although the LGM range of Atlantic cod was much smaller, and fragmented, both the ENMs agreed that populations should have been able to persist in suitable habitat on both sides of the Atlantic. The genetic results showed a degree of trans-Atlantic divergence consistent with genealogically continuous populations on both sides of the North Atlantic since long before the LGM, confirming the ENM results. In contrast, both the ENMs and the genetic data suggest that the Greenland G. morhua population post-dates the LGM.     

Modeling microalgae cultivation productivities in different geographic locations - estimation method for idealized photobioreactors

Microalgae can be used to produce versatile high-value fuels, such as methane, biodiesel, ethanol, or hydrogen gas. One of the most important factors that influence the economics of microalgae cultivation is the primary production of biomass per unit area. This is determined by productivity rates during cultivation, which are influenced by the local climate conditions (solar irradiation, temperature). To compare locations in different climate regions for microalgae cultivation, a mathematical model for an idealized closed photobioreactor was developed. The applied growth kinetics were based on theoretical maximum photon-conversion efficiencies (for the conversion of solar energy to chemical energy in the form of biomass). Known or estimated temperature effects for different algal strains were incorporated. The model was used to calculate hourly average areal productivity rates as well as annual primary production values under local conditions at seven example locations. Here, hourly weather data (solar irradiance and air temperature) were taken into account. According to these model calculations, maximum annual yields were achieved in regions with high irradiation and temperature patterns in or near the optimum range of the specific algal strain (here, desert and equatorial humid climates). The developed model can be used as a tool to assess and compare individual locations for microalgae cultivation.     

Mutation of albedo and growth response produces oscillations in a spatial Daisyworld

We extended a two-dimensional cellular automaton (CA) Daisyworld to include mutation of optimal growth temperature as well as mutation of albedo. Thus, the organisms (daisies) can adapt to prevailing environmental conditions or evolve to alter their environment. We find the resulting system oscillates with a period of hundreds of daisy generations. Weaker and less regular oscillations exist in previous daisyworld models, but they become much stronger and more regular here with mutation in the growth response. Despite the existence of a particular combination of mean albedo and optimum individual growth temperature which maximises growth, we find that this global state is unstable with respect to mutations which lower absolute growth rate, but increase marginal growth rate. The resulting system oscillates with a period that is found to decrease with increasing death rate, and to increase with increasing heat diffusion and heat capacity. We speculate that the origin of this oscillation is a Hopf bifurcation, previously predicted in a zero-dimensional system.     

Light Distribution and Photosynthesis in Field Crops

In a new model of light distribution in field crops a parameters is the fraction of light passing through unit leaf layer withoutinterception. Radiation profiles measured with solarimetersand photocells give values of s from 0.7 for grasses to 0.4for species with prostrate leaves. Knowing s, leaf transmissionT and leaf-area index L the light distribution in a field cropmay be described by a binomial expansion of the form {s+(I-s)T)^L.To calculate crop photosynthesis at given light intensity thisexpansion is combined with two parameters describing the shapeof the light-response curve of single leaves. Finally, the assumptionthat solar radiation varies sinusoidally allows daily totalphotosynthesis to be estimated from daylength and insolation. The theory predicts about the same potential photosynthesisin a cloudy temperte climate with long days as in a more sunnyequatorial climate with short days. When L < 3 photosynthesisincreases as s decreases, i.e. as leaves become more prostrate;but when L > 5, photosynthesis increases as s increases,i. e. as leaves become more erect. Assuming that respirationis proportional to leaf area, estimated dry-matter productionagrees well with field measurements on sugar-beet, sugar-cane,kale, and subterranean clover. Estimates of maximum gross photosynthesis(for sugar-cane and maize) range from 60 to 9 g m^–2 day^–1depending on insolation.     

干旱半干旱区灌丛斑块与降水量响应关系的熵模型模拟

灌丛斑块分布格局是灌木在干旱缺水条件下对生存环境的自我调节和适应的具体表现。应用熵理论和Klausmier模型,解释了灌丛斑块水分聚集原理并模拟了不同年降水条件下灌丛斑块的最佳面积比值(即最佳灌丛盖度)。研究结果表明:灌丛斑块生物量与其土壤含水量呈反比例函数关系,当生态系统处于稳定状态时(即熵最大状况下),年降水量与灌丛斑块面积比值符合一定的线性关系。研究采用内蒙古草原地区的野外调查数据,获得模型所需参数,进而模拟了不同年降水量条件下灌丛斑块最佳面积比值,研究结果可为半干旱地区植被保护与恢复提供参考。     

Hawaii: an ideal model for international aerobiological research

Future aerobiological studies important to mankind's understanding of our planet must demonstrate the ecological interrelationships between the aerial biota of island ecosystems and meteorological, chemical, physical and geographic factors. For example, air quality as related to human health and plant pathology can be more easily ascertained using oceanic islands such as Hawaii with the establishment of an international interdisciplinary aerobiological research team. This revised version was published online in June 2006 with corrections to the Cover Date.     

Daisyworld is Darwinian: constraints on adaptation are important for planetary self-regulation

The Daisyworld model demonstrates that self-regulation of the global environment can emerge from competition amongst types of life altering their local environment in different ways. Robertson & Robinson (1998. J. theor. Biol.195, 129-134) presented what they describe as a "Darwinian Daisyworld" in which the ability of organisms to adapt their internal physiology in response to environmental change undermines their ability to regulate their environment. They assume that there are no bounds on the environmental conditions that organisms can adapt to and that equal growth rates can potentially be achieved under any conditions. If adaptation could respond sufficiently rapidly to changes in the environment, this would eliminate any need for the environment to be regulated in the first place, because all possible states of the environment would be equally tolerable to life. However, the thermodynamics, chemistry and structure of living organisms set bounds on the range of environmental conditions that can be adapted to. As these bounds are approached, environmental conditions limit growth rate, and adaptations necessary for survival can also cost energy. Here we take account of such constraints and find that environmental regulation is recovered in the Daisyworld model. Hence, we suggest that constraints are an important part of a self-regulating planetary system.     

Contemporary climate change in the Sonoran Desert favors cold‐adapted species

Impacts of long‐term climate shifts on the dynamics of intact communities within species ranges are not well understood. Here, we show that warming and drying of the Southwestern United States over the last 25 years has corresponded to a shift in the species composition of Sonoran Desert winter annuals, paradoxically favoring species that germinate and grow best in cold temperatures. Winter rains have been arriving later in the season, during December rather than October, leading to the unexpected result that plants are germinating under colder temperatures, shifting community composition to favor slow growing, water‐use efficient, cold‐adapted species. Our results demonstrate how detailed ecophysiological knowledge of individual species, combined with long‐term demographic data, can reveal complex and sometimes unexpected shifts in community composition in response to climate change. Further, these results highlight the potentially overwhelming impact of changes in phenology on the response of biota to a changing climate.     

Leaf size, specific leaf area and microhabitat distribution of chaparral woody plants: contrasting patterns in species level and community level analyses

We examined variation in leaf size and specific leaf area (SLA) in relation to the distribution of 22 chaparral shrub species on small-scale gradients of aspect and elevation. Potential incident solar radiation (insolation) was estimated from a geographic information system to quantify microclimate affinities of these species across north- and south-facing slopes. At the community level, leaf size and SLA both declined with increasing insolation, based on average trait values for the species found in plots along the gradient. However, leaf size and SLA were not significantly correlated across species, suggesting that these two traits are decoupled and associated with different aspects of performance along this environmental gradient. For individual species, SLA was negatively correlated with species distributions along the insolation gradient, and was significantly lower in evergreen versus deciduous species. Leaf size exhibited a negative but non-significant trend in relation to insolation distribution of individual species. At the community level, variance in leaf size increased with increasing insolation. For individual species, there was a greater range of leaf size on south-facing slopes, while there was an absence of small-leaved species on north-facing slopes. These results demonstrate that analyses of plant functional traits along environmental gradients based on community level averages may obscure important aspects of trait variation and distribution among the constituent species.     

Fine scale endemism on coral reefs: archipelagic differentiation in turbinid gastropods

The perceived wide geographic range of organisms in the sea, facilitated by ready dispersal of waterborne dispersal stages, is a challenge for hypotheses of marine speciation but a boon to efforts of marine conservation. Wide species ranges are especially striking in the reef-rich Indo-west Pacific, the largest and most diverse marine biogeographic region, extending across half the planet. The insular marine biota of the tropical Pacific is characterized by wide-ranging species and provides the most striking examples of long distance dispersal, with endemism largely confined to the most remote island groups. Here we show that the gastropod Astralium "rhodostomum" has developed endemic clades on almost every Pacific archipelago sampled, a pattern unprecedented in marine biogeography, and reminiscent of the terrestrial biota of oceanic islands. Mitochondrial DNA sequences indicate that this species-complex is comprised of at least 30 geographically isolated clades, separated by as little as 180 km. Evidence suggests that such fine scale endemism and high diversity is not exceptional, but likely characterizes a substantial fraction of the reef biota. These results imply that (1) marine speciation can regularly occur over much finer spatial scales than generally accepted, (2) the diversity of coral reefs is even higher than suggested by morphology-based estimates, and (3) conservation efforts need to focus at the archipelagic level in the sea as on land.     

Climatic seasonality challenges the stability of microbial-driven deep soil carbon accumulation across China

Microbial residues contribute to the long-term stabilization of carbon in the entire soil profile, helping to regulate the climate of the planet; however, how sensitive these residues are to climatic seasonality remains virtually unknown, especially for deep soils across environmental gradients. Here, we investigated the changes of microbial residues along soil profiles (0–100 cm) from 44 typical ecosystems with a wide range of climates (~3100 km transects across China). Our results showed that microbial residues account for a larger portion of soil carbon in deeper (60–100 cm) vs. shallower (0–30 and 30–60 cm) soils. Moreover, we find that climate especially challenges the accumulation of microbial residues in deep soils, while soil properties and climate share their roles in controlling the residue accumulation in surface soils. Climatic seasonality, including positive correlations with summer precipitation and maximum monthly precipitation, as well as negative correlations with temperature annual range, are important factors explaining microbial residue accumulation in deep soils across China. In particular, summer precipitation is the key regulator of microbial-driven carbon stability in deep soils, which has 37.2% of relative independent effects on deep-soil microbial residue accumulation. Our work provides novel insights into the importance of climatic seasonality in driving the stabilization of microbial residues in deep soils, challenging the idea that deep soils as long-term carbon reservoirs can buffer climate change.