Identifying and quantifying planetary boundaries that must not be transgressed could help prevent human activities from causing unacceptable environmental change, argue Johan Rockstrom and colleagues.

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A safe operating space for humanity

Humans are altering the composition of biological communities through a variety of activities that increase rates of species invasions and species extinctions, at all scales, from local to global. These changes in components of the Earth's biodiversity cause concern for ethical and aesthetic reasons, but they also have a strong potential to alter ecosystem properties and the goods and services they provide to humanity. Ecological experiments, observations, and theoretical developments show that ecosystem properties depend greatly on biodiversity in terms of the functional characteristics of organisms present in the ecosystem and the distribution and abundance of those organisms over space and time. Species effects act in concert with the effects of climate, resource availability, and disturbance regimes in influencing ecosystem properties. Human activities can modify all of the above factors; here we focus on modification of these biotic controls. The scientific community has come to a broad consensus on many aspects of the re- lationship between biodiversity and ecosystem functioning, including many points relevant to management of ecosystems. Further progress will require integration of knowledge about biotic and abiotic controls on ecosystem properties, how ecological communities are struc- tured, and the forces driving species extinctions and invasions. To strengthen links to policy and management, we also need to integrate our ecological knowledge with understanding of the social and economic constraints of potential management practices. Understanding this complexity, while taking strong steps to minimize current losses of species, is necessary for responsible management of Earth's ecosystems and the diverse biota they contain.

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Effects of biodiversity on ecosystem functioning: a consensus of current knowledge

Anthropogenic pressures on the Earth System have reached a scale where abrupt global environmental change can no longer be excluded. We propose a new approach to global sustainability in which we define planetary boundaries within which we expect that humanity can operate safely. Transgressing one or more planetary boundaries may be deleterious or even catastrophic due to the risk of crossing thresholds that will trigger non-linear, abrupt environmental change within continental- to planetary-scale systems. We have identified nine planetary boundaries and, drawing upon current scientific understanding, we propose quantifications for seven of them. These seven are climate change (CO2 concentration in the atmosphere <350 ppm and/or a maximum change of +1 W m-2 in radiative forcing); ocean acidification (mean surface seawater saturation state with respect to aragonite ≥ 80% of pre-industrial levels); stratospheric ozone (<5% reduction in O3 concentration from pre-industrial level of 290 Dobson Units); biogeochemical nitrogen (N) cycle (limit industrial and agricultural fixation of N2 to 35 Tg N yr-1) and phosphorus (P) cycle (annual P inflow to oceans not to exceed 10 times the natural background weathering of P); global freshwater use (<4000 km3 yr-1 of consumptive use of runoff resources); land system change (<15% of the ice-free land surface under cropland); and the rate at which biological diversity is lost (annual rate of <10 extinctions per million species). The two additional planetary boundaries for which we have not yet been able to determine a boundary level are chemical pollution and atmospheric aerosol loading. We estimate that humanity has already transgressed three planetary boundaries: for climate change, rate of biodiversity loss, and changes to the global nitrogen cycle. Planetary boundaries are interdependent, because transgressing one may both shift the position of other boundaries or cause them to be transgressed. The social impacts of transgressing boundaries will be a function of the social-ecological resilience of the affected societies. Our proposed boundaries are rough, first estimates only, surrounded by large uncertainties and knowledge gaps. Filling these gaps will require major advancements in Earth System and resilience science. The proposed concept of "planetary boundaries" lays the groundwork for shifting our approach to governance and management, away from the essentially sectoral analyses of limits to growth aimed at minimizing negative externalities, toward the estimation of the safe space for human development. Planetary boundaries define, as it were, the boundaries of the "planetary playing field" for humanity if we want to be sure of avoiding major human-induced environmental change on a global scale.

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Planetary boundaries: Exploring the safe operating space for humanity

When a person looks at an object while exploring it with their hand, vision and touch both provide information for estimating the properties of the object. Vision frequently dominates the integrated visual-haptic percept, for example when judging size, shape or position, but in some circumstances the percept is clearly affected by haptics. Here we propose that a general principle, which minimizes variance in the final estimate, determines the degree to which vision or haptics dominates. This principle is realized by using maximum-likelihood estimation to combine the inputs. To investigate cue combination quantitatively, we first measured the variances associated with visual and haptic estimation of height. We then used these measurements to construct a maximum-likelihood integrator. This model behaved very similarly to humans in a visual-haptic task. Thus, the nervous system seems to combine visual and haptic information in a fashion that is similar to a maximum-likelihood integrator. Visual dominance occurs when the variance associated with visual estimation is lower than that associated with haptic estimation.

Humans integrate visual and haptic information in a statistically optimal fashion.

The ecological consequences of biodiversity loss have aroused considerable interest and controversy during the past decade. Major advances have been made in describing the relationship between species diversity and ecosystem processes, in identifying functionally important species, and in revealing underlying mechanisms. There is, however, uncertainty as to how results obtained in recent experiments scale up to landscape and regional levels and generalize across ecosystem types and processes. Larger numbers of species are probably needed to reduce temporal variability in ecosystem processes in changing environments. A major future challenge is to determine how biodiversity dynamics, ecosystem processes, and abiotic factors interact.

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Biodiversity and Ecosystem Functioning: Current Knowledge and Future Challenges

Most predictions of plant species effects on ecosystems are based on single traits (e.g., litter chemistry) or suites of related traits (functional groups). However, recent studies demonstrate that predictions of species effects on ecosystems are improved by considering multiple traits. In order to develop this multiple trait approach, it is critical to understand how these multiple traits vary in relation to one another among species. The ecosystem effects of traits that strongly covary can likely be summarized by one of these traits. In contrast, it will be necessary to determine the ecosystem effects of specific trait combinations for those traits that vary independently across species. In the field, I established monocultures of eight herbaceous species common in California annual grasslands. Plant species significantly differed in their litter quantity and quality, live biomass, and effects on soil labile C, soil temperature, and soil moisture. Species effects on soil moisture and temperature were only significant at the times of the growing season when each of these limited plant and microbial activity. Some of these traits cor- related with one another, such as litter biomass and species effects on soil temperature during the winter. However, for the most part, plant species exhibited unique combinations of these traits. For example, species with similar litter chemistry had the largest differences in plant biomass, soil moisture, and soil labile C. Species rankings for many traits changed over the growing season (e.g., biomass), so that the relationship among traits varied sea- sonally. The independent variation of these traits suggests that predictions of plant species effects on ecosystems will likely be enhanced by an understanding of how the ecosystem effects of plant traits may vary depending on the combination of traits.

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Plant traits that influence ecosystem processes vary independently among species

Effects of host diversity on disease dynamics.

The adenovirus E1A oncogene products stimulate DNA synthesis and cell proliferation but fail to transform primary baby rat kidney (BRK) cells because of the induction of p53-mediated programmed cell death (apoptosis). Overexpression of dominant mutant p53 (to abrogate wild-type p53 function) or introduction of apoptosis inhibitors, such as adenovirus E1B 19K or Bcl-2 oncoproteins, prevents E1A-induced apoptosis and permits transformation of BRK cells. The ability of activated Harvey-ras (H-ras) to cooperate with E1A to transform BRK cells suggests that H-ras is capable of overcoming the E1A-induced, p53-dependent apoptosis. We demonstrate here that activated H-ras was capable of suppressing apoptosis induced by E1A and wild-type p53. However, unlike Bcl-2 and the E1B 19K proteins, which completely block apoptosis but not p53-dependent growth arrest, H-ras expression permitted DNA synthesis and cell proliferation in the presence of high levels of wild-type p53. The mechanism by which H-ras regulates apoptosis and cell cycle progression is thereby strikingly different from that of the E1B 19K and Bcl-2 proteins. BRK cells transformed with H-ras and the temperature sensitive murine mutant p53(val 135), which lack E1A, underwent growth arrest at the permissive temperature for wild-type p53. p53-dependent growth arrest, however, could be relieved by E1A expression. Thus, H-ras alone was insufficient and cooperation of H-ras and E1A was required to override growth suppression by p53. Our data further suggest that two complementary growth signals from E1A plus H-ras can rescue cell death and thus permit transformation.

Activated H-ras rescues E1A-induced apoptosis and cooperates with E1A to overcome p53-dependent growth arrest.

Plant invasions can have large effects on ecosystem services. Some plant invaders were introduced specifically to restore key services to ecosystems, and other invaders are having unintended, detrimental effects on services, such as the quantity and quality of water delivered, flood control, erosion control, and food production. Many ecosystem services are difficult to measure directly, and although there are extensive studies on plant invaders and ecosystem processes, a number of challenges prevent us from confidently extrapolating those processes as proxies for services. To extrapolate local, short-term measures of processes to ecosystem services, we must: (1) determine which processes are the key contributors to a service, (2) assess how multiple processes interact to provide a given service, (3) determine how vegetation types and species affect those processes, and (4) explicitly assess how ecosystem services and their controls vary over space and time, including reliance of ecosystem service...

https://bioone.org/journals/invasive-plant-science-and-management/volume-5/issue-1/IPSM-D-11-00095.1/Measuring-the-Effects-of-Invasive-Plants-on-Ecosystem-Services/10.1614/IPSM-D-11-00095.1.pdf

Measuring the Effects of Invasive Plants on Ecosystem Services: Challenges and Prospects

Current knowledge of plant-soil feedback is based largely on single end point studies with soils conditioned by monocultures, but accounting for variability in the ecological impacts of feedback effects may require understanding how feedback develops over time and in multi-species plant communities. To examine temporal development and additivity of feedback, two pairs of native and non-native congeneric grasses were grown alone or in mixtures to create six soil conditioning treatments. We measured plant growth and feedback on the soils over 19 months and addressed whether plant biomass was additive or non-additive between soils treated by mixtures and their constituent monocultures. For native grasses, plant-soil feedback either became progressively more negative through time or switched from neutral to negative. Feedback to non-native grasses was variably neutral to positive. Final biomass of the grasses growing on soils conditioned by mixtures was generally an additive function of growth on soils conditioned by the component monocultures, except native grasses growing in soils conditioned by their own congener mixtures, which were non-additive. Temporal variation and non-additivity in feedback suggest that extrapolation to communities may be complex. More work is needed to assess the generality of temporal and scaling effects.

Valerie T. Eviner

Papers

Plant traits that influence ecosystem processes vary independently among species

Effects of host diversity on disease dynamics.

Activated H-ras rescues E1A-induced apoptosis and cooperates with E1A to overcome p53-dependent growth arrest.

Measuring the Effects of Invasive Plants on Ecosystem Services: Challenges and Prospects

The temporal development and additivity of plant-soil feedback in perennial grasses