Atmosphere, ecology and evolution: what drove the Miocene expansion of C4 grasslands?

Grasses using the C4 photosynthetic pathway dominate today's savanna ecosystems and account for ∼20% of terrestrial carbon fixation. However, this dominant status was reached only recently, during a period of C4 grassland expansion in the Late Miocene and Early Pliocene (4–8 Myr ago). Declining atmospheric CO2 has long been considered the key driver of this event, but new geological evidence casts doubt on the idea, forcing a reconsideration of the environmental cues for C4 plant success.Here, I evaluate the current hypotheses and debate in this field, beginning with a discussion of the role of CO2 in the evolutionary origins, rather than expansion, of C4 grasses. Atmospheric CO2 starvation is a plausible selection agent for the C4 pathway, but a time gap of around 10 Myr remains between major decreases in CO2 during the Oligocene, and the earliest current evidence of C4 plants.An emerging ecological perspective explains the Miocene expansion of C4 grasslands via changes in climatic seasonality and the occurrence of fire. However, the climatic drivers of this event are debated and may vary among geographical regions.Uncertainty in these areas could be reduced significantly by new directions in ecological research, especially the discovery that grass species richness along rainfall gradients shows contrasting patterns in different C4 clades. By re-evaluating a published data set, I show that increasing seasonality of rainfall is linked to changes in the relative abundance of the major C4 grass clades Paniceae and Andropogoneae. I propose that the explicit inclusion of these ecological patterns would significantly strengthen climate change hypotheses of Miocene C4 grassland expansion. Critically, they allow a new series of testable predictions to be made about the fossil record.Synthesis. This paper offers a novel framework for integrating modern ecological patterns into theories about the geological history of C4 plants

Environmental trade-offs of pig production systems under varied operational efficiencies

Production of pork, the most consumed meat globally, is estimated to emit 668 m tonnes CO2-eq of greenhouse gases each year. Amongst various production systems that comprise the pig industry, grainbased intensive production is widely regarded as the largest polluter of the environment, and thus it is imperative to develop alternative systems that can provide the right balance between sustainability and food security. Using an original dataset from the Republic of Ireland, this paper examines the life-cycle environmental impacts of representative pig farms operating under varying production efficiencies. For the baseline farm with an average production efficiency, global warming potential (GWP), acidifi- cation potential (AP) and eutrophication potential (EP) per kg carcass weight departing the slaughterhouse were estimated to be 3.5 kg CO2-eq, 43.8 g SO2-eq and 32.1 g PO4-eq, respectively. For herds with a higher production efficiency, a 9% improvement in feed conversion ratio was met by 6%, 15% and 12% decreases in GWP, EP, AP, respectively. Scenario and sensitivity analyses also revealed that (a) a switch to high-protein diets results in lower GWP and higher AP and EP, and (b) reducing transportation distances by sourcing domestically produced wheat and barley does not lower environmental impacts in any notable manner. To improve cross-study comparability of these findings, results based on an auxiliary functional unit, kg liveweight departing the farm gate, are also reported

Shared origins of a key enzyme during the evolution of C-4 and CAM metabolism

CAM and C4 photosynthesis are two key plant adaptations that have evolved independently multiple times, and are especially prevalent in particular groups of plants, including the Caryophyllales. We investigate the origin of photosynthetic PEPC, a key enzyme of both the CAM and C4 pathways. We combine phylogenetic analyses of genes encoding PEPC with analyses of RNA sequence data of Portulaca, the only plants known to perform both CAM and C4 photosynthesis. Three distinct gene lineages encoding PEPC exist in eudicots (namely ppc-1E1, ppc-1E2 and ppc-2), one of which (ppc-1E1) was recurrently recruited for use in both CAM and C4 photosynthesis within the Caryophyllales. This gene is present in multiple copies in the cacti and relatives, including Portulaca. The PEPC involved in the CAM and C4 cycles of Portulaca are encoded by closely related yet distinct genes. The CAM-specific gene is similar to genes from related CAM taxa, suggesting that CAM has evolved before C4 in these species. The similar origin of PEPC and other genes involved in the CAM and C4 cycles highlights the shared early steps of evolutionary trajectories towards CAM and C4, which probably diverged irreversibly only during the optimization of CAM and C4 phenotypes