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David J Beerling

Publications and source records attributed to David J Beerling.

9 recordsLinked to original sources

Zimmermann's telome theory of megaphyll leaf evolution: a molecular and cellular critique.

Megaphyll leaf evolution was a critical event in Earth history that had major consequences for the biotic regulation of the global environment. Zimmermann's telome theory has been widely accepted for over seventy years as the leading explanation for this evolutionary innovation. According to the telome theory, megaphylls evolved from the three-dimensional lateral branches of early vascular land plants in a hypothetical series of three transformations; first, the formation of determinate lateral branches (overtopping); second, the development of 'flattened' branch systems (planation); and third, the fusion of planated branches with lateral outgrowths of photosynthetic mesophyll tissue to form the leaf blade (webbing). A critical review of the molecular and cellular evidence identifies plausible genetic, cellular and physiological mechanisms in extant higher plants for overtopping and planation but more limited evidence for the process of webbing (lateral outgrowth fusion). We highlight key outstanding questions concerning the telome theory that are likely to be resolved when gene identification and functional analysis techniques are applied to photosynthetic organisms that have different evolutionary histories.

Biological Evolution↗

Nature's green revolution: the remarkable evolutionary rise of C4 plants.

Plants with the C4 photosynthetic pathway dominate today's tropical savannahs and grasslands, and account for some 30% of global terrestrial carbon fixation. Their success stems from a physiological CO2-concentrating pump, which leads to high photosynthetic efficiency in warm climates and low atmospheric CO2 concentrations. Remarkably, their dominance of tropical environments was achieved in only the past 10 million years (Myr), less than 3% of the time that terrestrial plants have existed on Earth. We critically review the proposal that declining atmospheric CO2 triggered this tropical revolution via its effects on the photosynthetic efficiency of leaves. Our synthesis of the latest geological evidence from South Asia and North America suggests that this emphasis is misplaced. Instead, we find important roles for regional climate change and fire in South Asia, but no obvious environmental trigger for C4 success in North America. CO2-starvation is implicated in the origins of C4 plants 25-32 Myr ago, raising the possibility that the pathway evolved under more extreme atmospheric conditions experienced 10 times earlier. However, our geochemical analyses provide no evidence of the C4 mechanism at this time, although possible ancestral components of the C4 pathway are identified in ancient plant lineages. We suggest that future research must redress the substantial imbalance between experimental investigations and analyses of the geological record.

Biological Evolution↗

Leaf evolution: gases, genes and geochemistry.

AIMS: This Botanical Briefing reviews how the integration of palaeontology, geochemistry and developmental biology is providing a new mechanistic framework for interpreting the 40- to 50-million-year gap between the origination of vascular land plants and the advent of large (megaphyll) leaves, a long-standing puzzle in evolutionary biology. SCOPE: Molecular genetics indicates that the developmental mechanisms required for leaf production in vascular plants were recruited long before the advent of large megaphylls. According to theory, this morphogenetic potential was only realized as the concentration of atmospheric CO2 declined during the late Palaeozoic. Surprisingly, plants effectively policed their own evolution since the decrease in CO2 was brought about as terrestrial floras evolved accelerating the rate of silicate rock weathering and enhancing sedimentary organic carbon burial, both of which are long-term sinks for CO2. CONCLUSIONS: The recognition that plant evolution responds to and influences CO(2) over millions of years reveals the existence of an intricate web of vegetation feedbacks regulating the long-term carbon cycle. Several of these feedbacks destabilized CO2 and climate during the late Palaeozoic but appear to have quickened the pace of terrestrial plant and animal evolution at that time.

Atmosphere↗

Feedbacks and the coevolution of plants and atmospheric CO2.

The coupled evolution of land plants, CO2, and climate over the last half billion years has maintained atmospheric CO2 concentrations within finite limits, indicating the involvement of a complex network of geophysiological feedbacks. But insight into this important regulatory network is extremely limited. Here we present a systems analysis of the physiological and geochemical processes involved, identifying new positive and negative feedbacks between plants and CO2 on geological time scales. Positive feedbacks accelerated falling CO2 concentrations during the evolution and diversification of terrestrial ecosystems in the Paleozoic and enhanced rising CO2 concentrations across the Triassic-Jurassic boundary during flood basalt eruptions. The existence of positive feedbacks reveals the unexpected destabilizing influence of the biota in climate regulation that led to environmental modifications accelerating rates of terrestrial plant and animal evolution in the Paleozoic.

Biological Evolution↗

A humid climate state during the Palaeocene/Eocene thermal maximum.

An abrupt climate warming of 5 to 10 degrees C during the Palaeocene/Eocene boundary thermal maximum (PETM) 55 Myr ago is linked to the catastrophic release of approximately 1,050-2,100 Gt of carbon from sea-floor methane hydrate reservoirs. Although atmospheric methane, and the carbon dioxide derived from its oxidation, probably contributed to PETM warming, neither the magnitude nor the timing of the climate change is consistent with direct greenhouse forcing by the carbon derived from methane hydrate. Here we demonstrate significant differences between marine and terrestrial carbon isotope records spanning the PETM. We use models of key carbon cycle processes to identify the cause of these differences. Our results provide evidence for a previously unrecognized discrete shift in the state of the climate system during the PETM, characterized by large increases in mid-latitude tropospheric humidity and enhanced cycling of carbon through terrestrial ecosystems. A more humid atmosphere helps to explain PETM temperatures, but the ultimate mechanisms underlying the shift remain unknown.

Atmosphere↗

Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum.

The aims of this paper are to review previously published palaeovegetation and independent palaeoclimatic datasets together with new results we present from dynamic vegetation model simulations and modern pollen rain studies to: (i) determine the responses of Amazonian ecosystems to changes in temperature, precipitation and atmospheric CO2 concentrations that occurred since the last glacial maximum (LGM), ca. 21 000 years ago; and (ii) use this long-term perspective to predict the likely vegetation responses to future climate change. Amazonia remained predominantly forested at the LGM, although the combination of reduced temperatures, precipitation and atmospheric CO2 concentrations resulted in forests structurally and floristically quite different from those of today. Cold-adapted Andean taxa mixed with rainforest taxa in central areas, while dry forest species and lianas probably became important in the more seasonal southern Amazon forests and savannahs expanded at forest-savannah ecotones. Net primary productivity (NPP) and canopy density were significantly lower than today. Evergreen rainforest distribution and NPP increased during the glacial-Holocene transition owing to ameliorating climatic and CO2 conditions. However, reduced precipitation in the Early-Mid-Holocene (ca. 8000-3600 years ago) caused widespread, frequent fires in seasonal southern Amazonia, causing increased abundance of drought-tolerant dry forest taxa and savannahs in ecotonal areas. Rainforests expanded once more in the Late Holocene owing to increased precipitation caused by greater austral summer insolation, although some of this forest expansion (e.g. in parts of the Bolivian Beni) is clearly caused by palaeo Indian landscape modification. The plant communities that existed during the Early-Mid-Holocene may provide insights into the kinds of vegetation response expected from similar increases in temperature and aridity predicted for the twenty-first century. We infer that ecotonal areas near the margins of the Amazon Basin are liable to be most sensitive to future environmental change and should therefore be targeted with conservation strategies that allow 'natural' species movements and plant community re-assortments to occur.

Atmosphere↗

The penalty of a long, hot summer. Photosynthetic acclimation to high CO2 and continuous light in "living fossil" conifers.

Deciduous forests covered the ice-free polar regions 280 to 40 million years ago under warm "greenhouse" climates and high atmospheric pCO2. Their deciduous habit is frequently interpreted as an adaptation for minimizing carbon losses during winter, but experiments with "living fossils" in a simulated warm polar environment refute this explanation. Measured carbon losses through leaf abscission of deciduous trees are significantly greater than losses through winter respiration in evergreens, yet annual rates of primary productivity are similar in all species. Here, we investigate mechanisms underlying this apparent paradox by measuring the seasonal patterns of leaf photosynthesis (A) under pCO2 enrichment in the same trees. During spring, A increased significantly in coastal redwood (Sequoia sempervirens), dawn redwood (Metasequoia glyptostroboides), and swamp cypress (Taxodium distichum) at an elevated pCO2 of 80 Pa compared with controls at 40 Pa. However, strong acclimation in Rubisco carboxylation capacity (Vc,max) completely offset the CO2 response of A in all species by the end of 6 weeks of continuous illumination in the simulated polar summer. Further measurements demonstrated the temporary nature of acclimation, with increases in Vc,max during autumn restoring the CO2 sensitivity of A. Contrary to expectations, the acclimation of Vc,max was not always accompanied by accumulation of leaf carbohydrates, but was associated with a decline in leaf nitrogen in summer, suggesting an alteration of the balance in plant sources and sinks for carbon and nitrogen. Preliminary calculations using A indicated that winter carbon losses through deciduous leaf abscission and respiration were recovered by 10 to 25 d of canopy carbon fixation during summer, thereby explaining the productivity paradox.

Acclimatization↗

Carbon loss by deciduous trees in a CO2-rich ancient polar environment.

Fossils demonstrate that deciduous forests covered the polar regions for much of the past 250 million years when the climate was warm and atmospheric CO2 high. But the evolutionary significance of their deciduous character has remained a matter of conjecture for almost a century. The leading hypothesis argues that it was an adaptation to photoperiod, allowing the avoidance of carbon losses by respiration from a canopy of leaves unable to photosynthesize in the darkness of warm polar winters. Here we test this proposal with experiments using 'living fossil' tree species grown in a simulated polar climate with and without CO2 enrichment. We show that the quantity of carbon lost annually by shedding a deciduous canopy is significantly greater than that lost by evergreen trees through wintertime respiration and leaf litter production, irrespective of growth CO2 concentration. Scaling up our experimental observations indicates that the greater expense of being deciduous persists in mature forests, even up to latitudes of 83 degrees N, where the duration of the polar winter exceeds five months. We therefore reject the carbon-loss hypothesis as an explanation for the deciduous nature of polar forests.

Antarctic Regions↗

Variation in Ginkgo biloba L. leaf characters across a climatic gradient in China.

Fossil leaves assigned to the genus Ginkgo are increasingly being used to reconstruct Mesozoic and Tertiary environments based on their stomatal and carbon isotopic characteristics. We sought to provide a more secure basis for understanding variations seen in the plant fossil record by determining the natural variability of these properties of sun and shade leaf morphotypes of Ginkgo biloba trees under the present atmospheric CO2 concentration and a range of contemporary climates in three Chinese locations (Lanzhou, Beijing, and Nanjing). Climate had no major effects on leaf stomatal index (proportion of leaf surface cells that are stomata) but did result in more variable stomatal densities. The effects of climate and leaf morphotype on stomatal index were rather conserved (<1%) and much less than the response of trees to recent CO2 increases. Leaf carbon isotope discrimination (delta) was highest for trees in Nanjing, which experience a warm, moist climate, whereas trees in the most arid site (Lanzhou) had the lowest delta values. Interestingly, the variation in delta shown by leaf populations of trees from China and the United Kingdom was very similar to that of fossil Ginkgo cuticles dating to the Mesozoic and Tertiary, which suggests to us that the physiology of leaf carbon uptake and regulation of water loss in Ginkgo has remained highly conserved despite the potential for evolutionary change over millions of years.

Biological Evolution↗