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.     

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.     

Maximization principles and daisyworld

We investigate whether the equilibrium time-averaged state of a self-organizing system with many internal degrees of freedom, 2D-daisyworld, can be described by optimizing a single quantity. Unlike physical systems where a principle of maximum energy production has been observed, daisyworld follows evolutionary dynamics rather than Hamiltonian dynamics. We find that this is sufficient to invalidate the maximum entropy production principle, finding instead a different principle, that the system self-organizes to a state which maximizes the amount of life.     

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.     

基于系统动力学的雏菊世界模型气候控制敏感性分析

Lovelock于20世纪70年代提出了著名的盖娅假说,认为地球是一个由生物和非生物环境组成的“超级有机体”,有能力调节自身的气候和化学组成,使之适合于生物的生存.盖娅假说遭到了生物学家,尤其是新达尔文主义者的强烈批评,认为它是目的论;随后,Lovelock通过雏菊世界模型论证了该假说.模型中的黑雏菊和白雏菊通过反射较少或较多的太阳光来调节地球的温度.基于雏菊世界模型,通过假设不同的初始条件——不同初始面积分配和不同的太阳光反射率组合,运用系统动力学对雏菊世界的温度调节能力进行了敏感性分析.结果证明,既使初始条件有差异,雏菊世界仍能够通过自我调节机制调节系统稳定时的星球温度,达到最适合生物生存的温度.雏菊世界模型对于初始面积分配、光照反射率组合条件不敏感.最后,雏菊世界模型的优点在于系统分析思想和以及对地球系统的简化,但是随着人类活动对地球系统影响的日益显著,建议模型应包含更多的环境、生物变量以及反馈关系和人类活动因素,将盖娅系统融入到自然-人类耦合的大系统中.     

Environmental regulation can arise under minimal assumptions

Models that demonstrate environmental regulation as a consequence of organism and environment coupling all require a number of core assumptions. Many previous models, such as Daisyworld, require that certain environment-altering traits have a selective advantage when those traits also contribute towards global regulation. We present a model that results in the regulation of a global environmental resource through niche construction without employing this and other common assumptions. There is no predetermined environmental optimum towards which regulation should proceed assumed or coded into the model. Nevertheless, polymorphic stable states that resist perturbation emerge from the simulated co-evolution of organisms and environment. In any single simulation a series of different stable states are realised, punctuated by rapid transitions. Regulation is achieved through two main subpopulations that are adapted to slightly different resource values, which force the environmental resource in opposing directions. This maintains the resource within a comparatively narrow band over a wide range of external perturbations. Population driven oscillations in the resource appear to be instrumental in protecting the regulation against mutations that would otherwise destroy it. Sensitivity analysis shows that the regulation is robust to mutation and to a wide range of parameter settings. Given the minimal assumptions employed, the results could reveal a mechanism capable of environmental regulation through the by-products of organisms.     

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.     

Scale dependence in desert plant diversity

The relationship between the number of species and the area sampled is one of the oldest and best-documented patterns in community ecology. An equation of the form S = cA ^z describing more precisely the species–area relation for plant species in smaller area is proposed as a result of intensive examination of species presence. Several study and field data from a wild range of plant and animal taxa suggest that the slope, z, of a graph of the logarithm of species richness against the logarithm of area is not a constant to the grassland or woodland community. We collected replicated and randomized plant data at 6 spatial scales from 1 m² to 1 km² in the desert region of northwest China to identify the scale dependence in desert plant biodiversity. The results showed that the slope of the log–log plot varied systematically with spatial scale. The value of z was high (0.37) at small scales from 1 to 10 m² and it decreased with increased spatial scale subsequently. When spatial scales varied from 1 m² to 1 km², the value of z varied from 0.37 to 0.035, suggesting that desert plant diversity has strongly scale-dependence at the small scales (less than 100 m²). The result is different from the research of grassland and woodland communities.     

The effect of <Emphasis Type="Italic">Cercidium praecox</Emphasis> and <Emphasis Type="Italic">Prosopis laevigata</Emphasis> on vertical distribution of soil free-living nematode communities in the Tehuacán Desert,Mexico

Vegetation cover is known to act as an abiotic mediator influencing the structure of soil fauna communities in arid and semi-arid ecosystems. The aim of the current research was to determine the spatial dispersion of the soil free-living nematode community under the canopy of Cercidium praecox and Prosopis laevigata during the rainy season. These shrubs are the dominant plant associations in the western part of the Tehuacán-Cuicatlán Valley in Mexico. Soil samples were taken from each 10-cm depth between 0 and 50 cm in August 2004. Our results demonstrated that the abundance and structure of the soil free-living nematode communities in the study area were strongly dependent on plant effects, specified by limited factors such as soil moisture and organic matter availability. The greatest degree of abundance of soil-free-living nematodes (88%) was found in the upper (0–10 cm) soil layer. Plant parasites were the most abundant trophic group under the two plants (58 and 36% under Parkinsonia (Cercidium) praecox and Prosopis laevigata, respectively), whereas omnivore-predators were the most dominant (96%) in inter-plant spaces. The fungivore/bacterivore (F/B) ratio was found to be the most useful tool of the ecological indices tested in the present study, reflecting the vertical distribution of the free-living nematode communities beneath different plant species in the different soil layers. The soil free-living nematode communities and their vertical distribution were found to be affected by plant ecophysiological adaptation, soil moisture, and the interaction between them.     

Decomposition along a rainfall gradient in the Judean desert,Israel

Summary The effect of a rainfall gradient, from a semiarid to extremely arid, on decomposition, were studied in the Judean desert, Israel. During the study period, the rainfall gradient obtained ranged from 308 mm to 24.4 mm. There was a annual mass loss of approximately 20% and 16% in the semi-arid and extremely arid regions, respectively. No significant correlation was found between the total rainfall and total mass losses. The data suggest that in an area where the conditions are not suitable for biological activity, the decomposition processes result from abiotic conditions, like temperature and radiation.Dedicated in memory of Prof. M. Evenari     

生物结皮对3种荒漠草本植物光合生理特性的影响

通过人工去除生物结皮实验,研究古尔班通古特沙漠生物结皮对3种荒漠草本植物生理特性的影响,根据生物结皮对土壤养分和水分的影响,综合分析生物结皮对3种草本植物光合生理特性的影响。结果表明,生物结皮对3种荒漠草本植物光合生理的影响基本一致,在植物生长早期,生物结皮区的尖喙牻牛儿苗（Erodium oxyrrhynchum M.Bieb.）、条叶庭荠（Alyssum linifolium Steph.ex Willd.）和琉苞菊（Hyalea pulchella（Ldb.） C.Koch）的叶片相对含水量（RWC）、净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（E）、水分利用效率（WUE）、PSⅡ最大光化学量子效率（F_v/F_m）、PSⅡ实际光化学效率（φ_PSⅡ）和叶绿素（Chl）含量均高于去除生物结皮区;但是,在植物生长后期,却是去除生物结皮区的各项生理指标高。生物结皮对3种荒漠草本植物光合生理过程的影响与其对植物生长的影响基本一致,这与土壤养分和水分状况的变化密切相关。     

Carbon flux dynamics and its environmental controlling factors in a desert steppe

Net ecosystem carbon dioxide exchange (NEE) above a desert steppe in Inner Mongolia was measured with eddy covariance technique in growing season of 2008. Environmental factors were also observed during the same period. Our results show that the minimum of growing season daily cumulative NEE appeared on August 26 (?1.6 g Cm^?2), while the maximum appeared on June 11 (0.9 g Cm^?2). Monthly cumulative NEE distributed in the growing season unevenly. The minimum of monthly cumulative NEE appeared in June (0.74 g Cm^?2); the maximum appeared in September (?14.94 g Cm^?2). The sum of the growing season NEE was ?45.5 g Cm^?2. The most important environmental factor affecting the half-hour average NEE was photosynthetically active radiation. The most important environmental factor affecting the daily cumulative NEE was 20–30 cm soil water content.     

几种荒漠植物地上生物量估算的初步研究

在调查植物样方的基础上,利用植株冠幅特征如冠幅长与宽、株高、基径、新生枝条数、总枝条数等作为变量建立了估算典型荒漠植物地上生物量的模型,利用植株冠幅长与宽所建拟合方程对估算琵琶柴灌丛生物量的精确度较高,经验证,预测值与实测值的相关系数为O．9989,相对误差在4．79％～10．12％之间利用植株株高、基径所建拟合方程对估算梭梭、多枝柽柳生物量计算值与实测值的相关系数在0.9418以上,显著性检验表明相关系数均为极显著经验证,预测值与实测值的相关系数分别为0．9902和0．9875．相对误差分别为6．87％～19．22％和7．49％～18．47％。研究表明,在大面积荒漠植物生物量研究中,利用植株冠幅特征作为变量来估算琵琶柴灌丛地上生物量方法简便可行,用来估算梭梭、多枝柽柳等灌木地上生物量时存在一定误差     

Carbon flux dynamics and its environmental controlling factors in a desert steppe

Net ecosystem carbon dioxide exchange (NEE) above a desert steppe in Inner Mongolia was measured with eddy covariance technique in growing season of 2008. Environmental factors were also observed during the same period. Our results show that the minimum of growing season daily cumulative NEE appeared on August 26 (?1.6 g Cm^?2), while the maximum appeared on June 11 (0.9 g Cm^?2). Monthly cumulative NEE distributed in the growing season unevenly. The minimum of monthly cumulative NEE appeared in June (0.74 g Cm^?2); the maximum appeared in September (?14.94 g Cm^?2). The sum of the growing season NEE was ?45.5 g Cm^?2. The most important environmental factor affecting the half-hour average NEE was photosynthetically active radiation. The most important environmental factor affecting the daily cumulative NEE was 20–30 cm soil water content.     

Leaf-twig carbon isotope ratio differences in photosynthetic-twig desert shrubs

Summary Carbon isotope ratios were determined for bulk tissues of both leaves and current season twigs of 29 species of Mohave Desert shrubs. Leaf and twig tissues were found to differ in their carbon isotope ratios, only in those species which had photosynthetic twigs. These data suggest that the twigs of these species operate at lower intercellular CO₂ values than leaves, an interpretation which is consistent with available gas-exchange data. An effect of microhabitat was also evident between the mean isotope ratios of leaves from wash versus slope habitats.This research has been supported by NSF Grant BSR-8410943 and by DOE Grant DE-FGO2-86ER60399.     

不同干扰下荒漠啮齿动物种群动态的空间同步性

荒漠啮齿动物是动物生态学研究中的重要类群,对于丰富种群和群落生态学理论具有重要意义.国外关于荒漠啮齿动物的研究较多,尤以北美的研究较为深入,涉及群落组成(Brown,1973)、生态位(Harris, 1984;Kenagy and Bartholomew, 1985)、物种共存(Brown and Munger,1985)、资源分享及微生境选择(Brown and Lieberman,1973;Price,1978)等方面;而国内关于荒漠啮齿动物研究相对较少.     

毛乌素沙地的生态背景及其草地建设的原则与优化模式

 本文全面分析了毛乌素沙地生态过渡带的自然环境、植被的现状和历史变迁过程,以定性和定量相结合的方法阐述了环境和植被之间的动态关系,在此基础上提出了该地区草地建设的原则及优化生态模式,其中包括：水分平衡原则,半固定沙丘持续发展原则,网带状种植原则,滩地草、农、林复合系统模式、软梁半人工草地复合系统模式和硬梁天然草地放牧系统模式。     

Decomposition rate of ocotillo (Fouquieria splendens) wood in the desert of southern California and its use in estimating adult survival by life-cycle graph analysis

Dead and downed ocotillos (Fouquieria splendens) are common in the Colorado Desert of southern California and pose a question concerning the life-span of adult plants that have been reported to be long-lived: if ocotillo is long-lived, decay rates must be very slow. Our study focused on decomposition and disintegration of above-ground ocotillo wood over 14 years to obtain a rate of change in dead mass. Disintegration rate was used in a new application of life-cycle graph analysis to calculate persistence time of carcasses, which in turn was used to estimate adult survival rates. We conclude that previous estimates of life spans may be too conservative and half-life of adult shrubs is over 200 years. Our approach to estimating adult survival from decay rates provides an independent test of survival transitions based on live plants for some species and so is an additional tool for developing life cycle models.     

Stem photosynthesis in Psorothamnus spinosus (smoke tree) in the Sonoran desert of California

Summary Stem photosynthetic responses to environmental parameters were investigated with Psorothamnus spinosus in the Sonoran Desert of California. Light saturation of stem photosynthesis was equal to maximum midday summer irradance (1600–2000 mol·m^-2·s^-1). The optimum temperature for stem photosynthesis was 39°C, and lower stem temperatures (27–35°C) caused significant decreases (up to 50%) in stem photosynthesis. Positive stem photosynthesis was maintained up to 51°C. Stem photosynthesis was relatively insensitive to increasing vpd up to 5 kPa; However, stem conductance decreased by 25% at a vpd of 5 kPa. At vpd greater than 5 kPa stem photosynthesis decreased relatively more than that of stem conductance causing a decrease in water use efficiency and an increase an intercellular carbon dioxide concentration. Maximum stem photosynthetic rates were low (6.2–10.6 mol·m^-2·s^-1) on a stem surface area, but, stem photosynthetic rates of young shoots were substantially higher (19.5–33.3 mol· m^-2·s^-1) on a projected area basis.Dedicated to the memory of Dr. W.H. Muller