Extreme climatic events: impacts of drought and high temperature on physiological processes in agronomically important plants

Climate models predict more frequent and more severe extreme events (e.g., heat waves, extended drought periods, flooding) in many regions for the next decades. The impact of adverse environmental conditions on crop plants is ecologically and economically relevant. This review is focused on drought and heat effects on physiological status and productivity of agronomically important plants. Stomatal opening represents an important regulatory mechanism during drought and heat stress since it influences simultaneously water loss via transpiration and CO2 diffusion into the leaf apoplast which further is utilized in photosynthesis. Along with the reversible short-term control of stomatal opening, stomata and leaf epidermis may produce waxy deposits and irreversibly down-regulate the stomatal conductance and non-stomatal transpiration. As a consequence photosynthesis will be negatively affected. Rubisco activase—a key enzyme in keeping the Calvin cycle functional—is heat-sensitive and may become a limiting factor at elevated temperature. The accumulated reactive oxygen species (ROS) during stress represent an additional challenge under unfavorable conditions. Drought and heat cause accumulation of free amino acids which are partially converted into compatible solutes such as proline. This is accompanied by lower rates of both nitrate reduction and de novo amino acid biosynthesis. Protective proteins (e.g., dehydrins, chaperones, antioxidant enzymes or the key enzyme for proline biosynthesis) play an important role in leaves and may be present at higher levels under water deprivation or high temperatures. On the whole plant level, effects on long-distance translocation of solutes via xylem and phloem and on leaf senescence (e.g., anticipated, accelerated or delayed senescence) are important. The factors mentioned above are relevant for the overall performance of crops under drought and heat and must be considered for genotype selection and breeding programs.

Millennial and sub-millennial scale climatic variations recorded in polar ice cores over the last glacial period

Since its discovery in Greenland ice cores, the millennial scale climatic variability of the last glacial period has been increasingly documented at all latitudes with studies focusing mainly on Marine Isotopic Stage 3 (MIS 3; 28–60 thousand of years before present, hereafter ka) and characterized by short Dansgaard-Oeschger (DO) events. Recent and new results obtained on the EPICA and NorthGRIP ice cores now precisely describe the rapid variations of Antarctic and Greenland temperature during MIS 5 (73.5–123 ka), a time period corresponding to relatively high sea level. The results display a succession of abrupt events associated with long Greenland InterStadial phases (GIS) enabling us to highlight a sub-millennial scale climatic variability depicted by (i) short-lived and abrupt warming events preceding some GIS (precursor-type events) and (ii) abrupt warming events at the end of some GIS (rebound-type events). The occurrence of these sub-millennial scale events is suggested to be driven by the insolation at high northern latitudes together with the internal forcing of ice sheets. Thanks to a recent NorthGRIP-EPICA Dronning Maud Land (EDML) common timescale over MIS 5, the bipolar sequence of climatic events can be established at millennial to sub-millennial timescale. This shows that for extraordinary long stadial durations the accompanying Antarctic warming amplitude cannot be described by a simple linear relationship between the two as expected from the bipolar seesaw concept. We also show that when ice sheets are extensive, Antarctica does not necessarily warm during the whole GS as the thermal bipolar seesaw model would predict, questioning the Greenland ice core temperature records as a proxy for AMOC changes throughout the glacial period.

The dating of dipterocarp tree rings: establishing a record of carbon cycling and climatic change in the tropics

In a first step to obtain a proxy record of past climatic events (including the El Ni (n) over tildeo-Southern Oscillation) in the normally aseasonal tropical environment of Sabah, a radial segment from a recently fallen dipterocarp (Shorea Superba) was radiocarbon dated and subjected to carbon isotope analysis. The high-precision radiocarbon results fell into the ambiguous modern plateau where several calibrated dates can exist for each sample. Dating was achieved by wiggle matching using a Bayesian approach to calibration. Using the defined growth characteristics of Shorea superba, probability density distributions were calculated and improbable dates rejected. It was found that the tree most likely started growing around AD 1660-1685. A total of 173 apparent growth increments were measured and, therefore, it could be determined that the tree formed one ring approximately every two years. Stable carbon isotope values were obtained from resin-extracted wholewood from each ring. Carbon cycling is evident in the `juvenile effect', resulting from the assimilation of respired carbon dioxide and lower light levels below the canopy, and in the `anthropogenic effect' caused by increased industrial activity in the late-nineteenth and twentieth centuries. This study demonstrates that palaeoenvironmental information can be obtained from trees growing in aseasonal environments, where climatic conditions prevent the formation of well-defined annual rings.

A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphy

Due to their outstanding resolution and well-constrained chronologies, Greenland ice-core records provide a master record of past climatic changes throughout the Last Interglacial–Glacial cycle in the North Atlantic region. As part of the INTIMATE (INTegration of Ice-core, MArine and TErrestrial records) project, protocols have been proposed to ensure consistent and robust correlation between different records of past climate. A key element of these protocols has been the formal definition and ordinal numbering of the sequence of Greenland Stadials (GS) and Greenland Interstadials (GI) within the most recent glacial period. The GS and GI periods are the Greenland expressions of the characteristic Dansgaard–Oeschger events that represent cold and warm phases of the North Atlantic region, respectively. We present here a more detailed and extended GS/GI template for the whole of the Last Glacial period. It is based on a synchronization of the NGRIP, GRIP, and GISP2 ice-core records that allows the parallel analysis of all three records on a common time scale. The boundaries of the GS and GI periods are defined based on a combination of stable-oxygen isotope ratios of the ice (δ18O, reflecting mainly local temperature) and calcium ion concentrations (reflecting mainly atmospheric dust loading) measured in the ice. The data not only resolve the well-known sequence of Dansgaard–Oeschger events that were first defined and numbered in the ice-core records more than two decades ago, but also better resolve a number of short-lived climatic oscillations, some defined here for the first time. Using this revised scheme, we propose a consistent approach for discriminating and naming all the significant abrupt climatic events of the Last Glacial period that are represented in the Greenland ice records. The final product constitutes an extended and better resolved Greenland stratotype sequence, against which other proxy records can be compared and correlated. It also provides a more secure basis for investigating the dynamics and fundamental causes of these climatic perturbations.

Synchronous Holocene climatic oscillations recorded on the Swiss Plateau and at timberline in the Alps

Eight synchronous pre-Roman cold phases were found at 9600–9200, 8600–8150, 7550–6900, 6600– 6200, 5350–4900, 4600–4400, 3500–3200 and 2600–2350 radiocarbon years BP by reconstructing past climate at two sites on the Swiss Plateau and at timberline in the Alps. The cooling events during the early-and mid-Holocene represent temperature values similar to today, and apparently the onset of cooling events represents a deviation from today's mean annual temperature of about 1°C and is triggered at a 1000-year periodicity. At Wallisellen-Langachermoos (440 m), a former oligotrophic lake near Zürich, the correlation between sum mertime lake levels and the seed production of the amphi-Atlantic aquatic plantNajas flexilis was used to reconstruct lake levels over a 3000-year period during the first part of the Holocene. At Lake Seedorf on the western Swiss Plateau (609 m) the sedimentological, palynological and macrofossil record revealed fluctuations of lake levels for the complete Holocene. From Lago Basso in the southern Alps (2250 m, Val San Giacomo near Splügen Pass, Northern Italy) the terrestrial plant macrofossils – especiallyPinus cembra andLarix – allowed the reconstruction of timberline fluctuations controlled by climate. A similar climatic pattern was found at Gouillé Rion pond in the central Swiss Alps (2343 m, Val d'Hérémence) with plant macrofossils and pollen concentrations and percentages. We postulate that these climatic events are detectable throughout central Europe by independent methods in combination with precise AMS-radiocarbon datings on terrestrial plant remains. Our data fit other proxy records of regional climatic change, such as cool intervals from Greenland ice cores, glacier movements in the Swiss and Austrian Alps, and dendro-densitometry on subfossil wood, as well as the palaeoclimatic data from the Jura Mountains of France obtained by sedimentological analyses. Thus our data indicate that the Northern Hemisphere climate was less stable during the Holocene than previously believed.

Spatial and temporal variation in population genetic structure of wild Nile tilapia (Oreochromis niloticus) across Africa

Background: Reconstructing the evolutionary history of a species is challenging. It often depends not only on the past biogeographic and climatic events but also the contemporary and ecological factors, such as current connectivity and habitat heterogeneity. In fact, these factors might interact with each other and shape the current species distribution. However, to what extent the current population genetic structure reflects the past and the contemporary factors is largely unknown. Here we investigated spatio-temporal genetic structures of Nile tilapia (Oreochromis niloticus) populations, across their natural distribution in Africa. While its large biogeographic distribution can cause genetic differentiation at the paleo-biogeographic scales, its restricted dispersal capacity might induce a strong genetic structure at micro-geographic scales. Results: Using nine microsatellite loci and 350 samples from ten natural populations, we found the highest genetic differentiation among the three ichthyofaunal provinces and regions (Ethiopian, Nilotic and Sudano-Sahelian) (R(ST) = 0.38 - 0.69). This result suggests the predominant effect of paleo-geographic events at macro-geographic scale. In addition, intermediate divergences were found between rivers and lakes within the regions, presumably reflecting relatively recent interruptions of gene flow between hydrographic basins (R(ST) = 0.24 - 0.32). The lowest differentiations were observed among connected populations within a basin (R(ST) = 0.015 in the Volta basin). Comparison of temporal sample series revealed subtle changes in the gene pools in a few generations (F = 0 - 0.053). The estimated effective population sizes were 23 - 143 and the estimated migration rate was moderate (m similar to 0.094 - 0.097) in the Volta populations. Conclusions: This study revealed clear hierarchical patterns of the population genetic structuring of O. niloticus in Africa. The effects of paleo-geographic and climatic events were predominant at macro-geographic scale, and the significant effect of geographic connectivity was detected at micro-geographic scale. The estimated effective population size, the moderate level of dispersal and the rapid temporal change in genetic composition might reflect a potential effect of life history strategy on population dynamics. This hypothesis deserves further investigation. The dynamic pattern revealed at micro-geographic and temporal scales appears important from a genetic resource management as well as from a biodiversity conservation point of view.

Transient dominance in a central African rain forest

The large-crowned emergent tree Microberlinia bisulcata dominates rain forest groves at Korup National Park, Cameroon, along with two codominants, Tetraberlinia bifoliolata and T. korupensis. M. bisulcata has a pronounced modal size frequency distribution around 110 cm stem diameter: its recruitment potential is very poor. It is a long-lived light-demanding species, one of many found in African forests. Tetraberlinia species lack modality, are more shade tolerant, and recruit better. All three species are ectomycorrhizal. M. bisulcata dominates grove basal area, even though it has similar numbers of trees (≥50 cm stem diameter) as each of the other two species. This situation presented a conundrum that prompted a long-term study of grove dynamics. Enumerations of two plots (82.5 and 56.25 ha) between 1990 and 2010 showed mortality and recruitment of M. bisulcata to be very low (both rates 0.2% per year) compared with Tetraberlinia (2.4% and 0.8% per year), and M. bisulcata grows twice as fast as the Tetraberlinia. Ordinations indicated that these three species determined community structure by their strong negative associations while other species showed almost none. Ranked species abundance curves fitted the Zipf-Mandelbrot model well and allowed “overdominance” of M. bisulcata to be estimated. Spatial analysis indicated strong repulsion by clusters of large (50 to <100 cm) and very large (≥100 cm) M. bisulcata of their own medium-sized (10 to <50 cm) trees and all sizes of Tetraberlinia. This was interpreted as competition by M. bisulcata increasing its dominance, but also inhibition of its own replacement potential. Stem coring showed a modal age of 200 years for M. bisulcata, but with large size variation (50–150 cm). Fifty-year model projections suggested little change in medium, decreases in large, and increases in very large trees of M. bisulcata, accompanied by overall decreases in medium and large trees of Tetraberlinia species. Realistically increasing very-large-tree mortality led to grove collapse without short-term replacement. M. bisulcata most likely depends on climatic events to rebuild its stands: the ratio of disturbance interval to median species' longevity is important. A new theory of transient dominance explains how M. bisulcata may be cycling in abundance over time and displaying nonequilibrium dynamics.

North-south palaeohydrological contrasts in the central Mediterranean during the Holocene: tentative synthesis and working hypotheses

On the basis of a multi-proxy approach and a strategy combining lacustrine and marine records along a north–south transect, data collected in the central Mediterranean within the framework of a collaborative project have led to reconstruction of high-resolution and well-dated palaeohydrological records and to assessment of their spatial and temporal coherency. Contrasting patterns of palaeohydrological changes have been evidenced in the central Mediterranean: south (north) of around 40° N of latitude, the middle part of the Holocene was characterised by lake-level maxima (minima), during an interval dated to ca. 10 300–4500 cal BP to the south and 9000–4500 cal BP to the north. Available data suggest that these contrasting palaeohydrological patterns operated throughout the Holocene, both on millennial and centennial scales. Regarding precipitation seasonality, maximum humidity in the central Mediterranean during the middle part of the Holocene was characterised by humid winters and dry summers north of ca. 40° N, and humid winters and summers south of ca. 40° N. This may explain an apparent conflict between palaeoclimatic records depending on the proxies used for reconstruction as well as the synchronous expansion of tree species taxa with contrasting climatic requirements. In addition, south of ca. 40° N, the first millennium of the Holocene was characterised by very dry climatic conditions not only in the eastern, but also in the central- and the western Mediterranean zones as reflected by low lake levels and delayed reforestation. These results suggest that, in addition to the influence of the Nile discharge reinforced by the African monsoon, the deposition of Sapropel 1 has been favoured (1) by an increase in winter precipitation in the northern Mediterranean borderlands, and (2) by an increase in winter and summer precipitation in the southern Mediterranean area. The climate reversal following the Holocene climate optimum appears to have been punctuated by two major climate changes around 7500 and 4500 cal BP. In the central Mediterranean, the Holocene palaeohydrological changes developed in response to a combination of orbital, ice-sheet and solar forcing factors. The maximum humidity interval in the south-central Mediterranean started ca. 10 300 cal BP, in correlation with the decline (1) of the possible blocking effects of the North Atlantic anticyclone linked to maximum insolation, and/or (2) of the influence of the remnant ice sheets and fresh water forcing in the North Atlantic Ocean. In the north-central Mediterranean, the lake-level minimum interval began only around 9000 cal BP when the Fennoscandian ice sheet disappeared and a prevailing positive NAO-(North Atlantic Oscillation) type circulation developed in the North Atlantic area. The major palaeohydrological oscillation around 4500–4000 cal BP may be a non-linear response to the gradual decrease in insolation, with additional key seasonal and interhemispheric changes. On a centennial scale, the successive climatic events which punctuated the entire Holocene in the central Mediterranean coincided with cooling events associated with deglacial outbursts in the North Atlantic area and decreases in solar activity during the interval 11 700–7000 cal BP, and to a possible combination of NAO-type circulation and solar forcing since ca. 7000 cal BP onwards. Thus, regarding the centennial-scale climatic oscillations, the Mediterranean Basin appears to have been strongly linked to the North Atlantic area and affected by solar activity over the entire Holocene. In addition to model experiments, a better understanding of forcing factors and past atmospheric circulation patterns behind the Holocene palaeohydrological changes in the Mediterranean area will require further investigation to establish additional high-resolution and well-dated records in selected locations around the Mediterranean Basin and in adjacent regions. Special attention should be paid to greater precision in the reconstruction, on millennial and centennial timescales, of changes in the latitudinal location of the limit between the northern and southern palaeohydrological Mediterranean sectors, depending on (1) the intensity and/or characteristics of climatic periods/oscillations (e.g. Holocene thermal maximum versus Neoglacial, as well as, for instance, the 8.2 ka event versus the 4 ka event or the Little Ice Age); and (2) on varying geographical conditions from the western to the eastern Mediterranean areas (longitudinal gradients). Finally, on the basis of projects using strategically located study sites, there is a need to explore possible influences of other general atmospheric circulation patterns than NAO, such as the East Atlantic–West Russian or North Sea–Caspian patterns, in explaining the apparent complexity of palaeoclimatic (palaeohydrological) Holocene records from the Mediterranean area.

Promoting Local Innovations: A tool for adaptation to climate change and sustainable development in Mountains

Throughout their history mountain communities have had to adapt to changing environmental and socio-economic conditions. They have developed strategies and specialized knowledge to sustain their livelihoods in a context of adverse climatic events and constant change. As negotiations and discussions on climate change emphasize the critical need for locally relevant and community owned adaptation strategies, there is a need for new tools to capitalize on this local knowledge and endogenous potential for innovation. The toolkit Promoting Local Innovation (PLI) was designed by the Centre for Development and Environment (CDE) of the University of Bern, Switzerland, to facilitate a participatory social learning process which identifies locally available innovations that can be implemented for community development. It is based on interactive pedagogy and joint learning among different stakeholders in the local context. The tried-and-tested tool was developed in the Andean region in 2004, and then used in International Union for Conservation of Nature (IUCN) climate change adaptation projects in Thailand, Burkina Faso, Senegal, and Chile. These experiences showed that PLI can be used to involve all relevant stakeholders in establishing strategies and actions needed for rural communities to adapt to climate change impacts, while building on local innovation potential and promoting local ownership

Macroevolutionary patterns in the diversification of parrots: effects of climate change, geological events and key innovations

Aim Parrots are thought to have originated on Gondwana during the Cretaceous. The initial split within crown group parrots separated the New Zealand taxa from the remaining extant species and was considered to coincide with the separation of New Zealand from Gondwana 82-85 Ma, assuming that the diversification of parrots was mainly shaped by vicariance. However, the distribution patterns of several extant parrot groups cannot be explained without invoking transoceanic dispersal, challenging this assumption. Here, we present a temporal and spatial framework for the diversification of parrots using external avian fossils as calibration points in order to evaluate the relative importance of the influences of past climate change, plate tectonics and ecological opportunity. Location Australasian, African, Indo-Malayan and Neotropical regions. Methods Phylogenetic relationships were investigated using partial sequences of the nuclear genes c-mos, RAG-1 and Zenk of 75 parrot and 21 other avian taxa. Divergence dates and confidence intervals were estimated using a Bayesian relaxed molecular clock approach. Biogeographic patterns were evaluated taking temporal connectivity between areas into account. We tested whether diversification remained constant over time and if some parrot groups were more species-rich than expected given their age. Results Crown group diversification of parrots started only about 58 Ma, in the Palaeogene, significantly later than previously thought. The Australasian lories and possibly also the Neotropical Arini were found to be unexpectedly species-rich. Diversification rates probably increased around the Eocene/Oligocene boundary and in the middle Miocene, during two periods of major global climatic aberrations characterized by global cooling. Main conclusions The diversification of parrots was shaped by climatic and geological events as well as by key innovations. Initial vicariance events caused by continental break-up were followed by transoceanic dispersal and local radiations. Habitat shifts caused by climate change and mountain orogenesis may have acted as a catalyst to the diversification by providing new ecological opportunities and challenges as well as by causing isolation as a result of habitat fragmentation. The lories constitute the only highly nectarivorous parrot clade, and their diet shift, associated with morphological innovation, may have acted as an evolutionary key innovation, allowing them to explore underutilized niches and promoting their diversification.

Were last glacial climate events simultaneous between Greenland and western Europe?

During the last glacial period, several large abrupt climate fluctuations took place on the Greenland ice cap and elsewhere. Often these Dansgaard/Oeschger events are assumed to have been synchronous, and then used as tie-points to link chronologies between the proxy archives. However, if temporally separate events are lumped into one illusionary event, climatic interpretations of the tuned events will obviously be flawed. Here, we compare Dansgaard/Oeschger-type events in a well-dated record from south-eastern France with those in Greenland ice cores. Instead of assuming simultaneous climate events between both archives, we keep their age models independent. Even these well-dated archives possess large chronological uncertainties, that prevent us from inferring synchronous climate events at decadal to multi-centennial time scales. If possible, tuning of proxy archives should be avoided.

Relaxation of species-specific neighborhood effects in Bornean rain forest under climatic perturbation

Evidence of negative conspecific density dependence (NDD) operating on seedling survival and sapling recruitment has accumulated recently. In contrast, evidence of NDD operating on growth of trees has been circumstantial at best. Whether or not local NDD at the level of individual trees leads to NDD at the level of the community is still an open question. Moreover, whether and how perturbations interfere with these processes have rarely been investigated. We applied neighborhood models to permanent plot data from a Bornean dipterocarp forest censused over two 10-11 year periods. Although the first period was only lightly perturbed, a moderately strong El Nino event causing severe drought occurred in the first half of the second period. Such events are an important component of the environmental stochasticity affecting the region. We show that local NDD on growth of small-to-medium-sized trees may indeed translate to NDD at the level of the community. This interpretation is based on increasingly negative effects of bigger conspecific neighbors on absolute growth rates of individual trees with increasing basal area across the 18 most abundant overstory species in the first period. However, this relationship was much weaker in the second period. We interpreted this relaxation of local and community-level NDD as a consequence of increased light levels at the forest floor due to temporary leaf and twig loss of large trees in response to the drought event. Mitigation of NDD under climatic perturbation acts to decrease species richness, especially in forest overstory and therefore has an important role in determining species relative abundances at the site.

A tephra lattice for Greenland and a reconstruction of volcanic events spanning 25–45 ka b2k

Tephra layers preserved within the Greenland ice-cores are crucial for the independent synchronisation of these high-resolution records to other palaeoclimatic archives. Here we present a new and detailed tephrochronological framework for the time period 25,000 e 45,000 a b2k that brings together results from 4 deep Greenland ice-cores. In total, 99 tephra deposits, the majority of which are preserved as cryptotephra, are described from the NGRIP, NEEM, GRIP and DYE-3 records. The major element signatures of single glass shards within these deposits indicate that 93 are basaltic in composition all originating from Iceland. Specifically, 43 originate from Grimsv € otn, 20 are thought to be sourced from the Katla volcanic system and 17 show affinity to the Kverkfj € oll system. Robust geochemical characterisations, independent ages derived from the GICC05 ice-core chronology, and the stratigraphic positions of these deposits relative to the Dansgaard-Oeschger climate events represent a key framework that provides new information on the frequency and nature of volcanic events in the North Atlantic region between GS-3 and GI-12. Of particular importance are 19 tephra deposits that lie on the rapid climatic transitions that punctuate the last glacial period. This framework of well-constrained, time-synchronous tie-lines represents an important step towards the independent synchronisation of marine, terrestrial and ice-core records from the North Atlantic region, in order to assess the phasing of rapid climatic changes during the last glacial period.