Complex spatiotemporal haemodynamic response following sensory stimulation in the awake rat

Detailed understanding of the haemodynamic changes that underlie non-invasive neuroimaging techniques such as blood oxygen level dependent functional magnetic resonance imaging is essential if we are to continue to extend the use of these methods for understanding brain function and dysfunction. The use of animal and in particular rodent research models has been central to these endeavours as they allow in-vivo experimental techniques that provide measurements of the haemodynamic response function at high temporal and spatial resolution. A limitation of most of this research is the use of anaesthetic agents which may disrupt or mask important features of neurovascular coupling or the haemodynamic response function. In this study we therefore measured spatiotemporal cortical haemodynamic responses to somatosensory stimulation in awake rats using optical imaging spectroscopy. Trained, restrained animals received non-noxious stimulation of the whisker pad via chronically implanted stimulating microwires whilst optical recordings were made from the contralateral somatosensory cortex through a thin cranial window. The responses we measure from un-anaesthetised animals are substantially different from those reported in previous studies which have used anaesthetised animals. These differences include biphasic response regions (initial increases in blood volume and oxygenation followed by subsequent decreases) as well as oscillations in the response time series of awake animals. These haemodynamic response features do not reflect concomitant changes in the underlying neuronal activity and therefore reflect neurovascular or cerebrovascular processes. These hitherto unreported hyperemic response dynamics may have important implications for the use of anaesthetised animal models for research into the haemodynamic response function.

A three-compartment model of the hemodynamic response and oxygen delivery to brain

We describe a mathematical model linking changes in cerebral blood flow, blood volume and the blood oxygenation state in response to stimulation. The model has three compartments to take into account the fact that the cerebral blood flow and volume as measured concurrently using laser Doppler flowmetry and optical imaging spectroscopy have contributions from the arterial, capillary as well as the venous compartments of the vasculature. It is an extension to previous one-compartment hemodynamic models which assume that the measured blood volume changes are from the venous compartment only. An important assumption of the model is that the tissue oxygen concentration is a time varying state variable of the system and is driven by the changes in metabolic demand resulting from changes in neural activity. The model takes into account the pre-capillary oxygen diffusion by flexibly allowing the saturation of the arterial compartment to be less than unity. Simulations are used to explore the sensitivity of the model and to optimise the parameters for experimental data. We conclude that the three-compartment model was better than the one-compartment model at capturing the hemodynamics of the response to changes in neural activation following stimulation.

A model of the hemodynamic response and oxygen delivery to brain

A recent nonlinear system by Friston et al. (2000. NeuroImage 12: 466–477) links the changes in BOLD response to changes in neural activity. The system consists of five subsystems, linking: (1) neural activity to flow changes; (2) flow changes to oxygen delivery to tissue; (3) flow changes to changes in blood volume and venous outflow; (4) changes in flow, volume, and oxygen extraction fraction to deoxyhemoglobin changes; and finally (5) volume and deoxyhemoglobin changes to the BOLD response. Friston et al. exploit, in subsystem 2, a model by Buxton and Frank coupling flow changes to changes in oxygen metabolism which assumes tissue oxygen concentration to be close to zero. We describe below a model of the coupling between flow and oxygen delivery which takes into account the modulatory effect of changes in tissue oxygen concentration. The major development has been to extend the original Buxton and Frank model for oxygen transport to a full dynamic capillary model making the model applicable to both transient and steady state conditions. Furthermore our modification enables us to determine the time series of CMRO2 changes under different conditions, including CO2 challenges. We compare the differences in the performance of the “Friston system” using the original model of Buxton and Frank and that of our model. We also compare the data predicted by our model (with appropriate parameters) to data from a series of OIS studies. The qualitative differences in the behaviour of the models are exposed by different experimental simulations and by comparison with the results of OIS data from brief and extended stimulation protocols and from experiments using hypercapnia.

Equator-to-pole temperature differences and the extra-tropical storm track responses of the CMIP5 climate models

This paper aims to understand the physical processes causing the large spread in the storm track projections of the CMIP5 climate models. In particular, the relationship between the climate change responses of the storm tracks, as measured by the 2–6 day mean sea level pressure variance, and the equator-to-pole temperature differences at upper- and lower-tropospheric levels is investigated. In the southern hemisphere the responses of the upper- and lower-tropospheric temperature differences are correlated across the models and as a result they share similar associations with the storm track responses. There are large regions in which the storm track responses are correlated with the temperature difference responses, and a simple linear regression model based on the temperature differences at either level captures the spatial pattern of the mean storm track response as well explaining between 30 and 60 % of the inter-model variance of the storm track responses. In the northern hemisphere the responses of the two temperature differences are not significantly correlated and their associations with the storm track responses are more complicated. In summer, the responses of the lower-tropospheric temperature differences dominate the inter-model spread of the storm track responses. In winter, the responses of the upper- and lower-temperature differences both play a role. The results suggest that there is potential to reduce the spread in storm track responses by constraining the relative magnitudes of the warming in the tropical and polar regions.

On the lack of stratospheric dynamical variability in low-top versions of the CMIP5 models

We describe the main differences in simulations of stratospheric climate and variability by models within the fifth Coupled Model Intercomparison Project (CMIP5) that have a model top above the stratopause and relatively fine stratospheric vertical resolution (high-top), and those that have a model top below the stratopause (low-top). Although the simulation of mean stratospheric climate by the two model ensembles is similar, the low-top model ensemble has very weak stratospheric variability on daily and interannual time scales. The frequency of major sudden stratospheric warming events is strongly underestimated by the low-top models with less than half the frequency of events observed in the reanalysis data and high-top models. The lack of stratospheric variability in the low-top models affects their stratosphere-troposphere coupling, resulting in short-lived anomalies in the Northern Annular Mode, which do not produce long-lasting tropospheric impacts, as seen in observations. The lack of stratospheric variability, however, does not appear to have any impact on the ability of the low-top models to reproduce past stratospheric temperature trends. We find little improvement in the simulation of decadal variability for the high-top models compared to the low-top, which is likely related to the fact that neither ensemble produces a realistic dynamical response to volcanic eruptions.

A multimodel assessment of future projections of North Atlantic and European extratropical cyclones in the CMIP5 climate models

The response of North Atlantic and European extratropical cyclones to climate change is investigated in the climate models participating in phase 5 of the Coupled Model Intercomparison Project (CMIP5). In contrast to previous multimodel studies, a feature-tracking algorithm is here applied to separately quantify the re- sponses in the number, the wind intensity, and the precipitation intensity of extratropical cyclones. Moreover, a statistical framework is employed to formally assess the uncertainties in the multimodel projections. Under the midrange representative concentration pathway (RCP4.5) emission scenario, the December–February (DJF) response is characterized by a tripolar pattern over Europe, with an increase in the number of cyclones in central Europe and a decreased number in the Norwegian and Mediterranean Seas. The June–August (JJA) response is characterized by a reduction in the number of North Atlantic cyclones along the southern flank of the storm track. The total number of cyclones decreases in both DJF (24%) and JJA (22%). Classifying cyclones according to their intensity indicates a slight basinwide reduction in the number of cy- clones associated with strong winds, but an increase in those associated with strong precipitation. However, in DJF, a slight increase in the number and intensity of cyclones associated with strong wind speeds is found over the United Kingdom and central Europe. The results are confirmed under the high-emission RCP8.5 scenario, where the signals tend to be larger. The sources of uncertainty in these projections are discussed.

A GCSS model intercomparison for a tropical squall line observed during toga-coare. II: Intercomparison of single-column models and a cloud-resolving model

This paper presents single-column model (SCM) simulations of a tropical squall-line case observed during the Coupled Ocean-Atmosphere Response Experiment of the Tropical Ocean/Global Atmosphere Programme. This case-study was part of an international model intercomparison project organized by Working Group 4 ‘Precipitating Convective Cloud Systems’ of the GEWEX (Global Energy and Water-cycle Experiment) Cloud System Study. Eight SCM groups using different deep-convection parametrizations participated in this project. The SCMs were forced by temperature and moisture tendencies that had been computed from a reference cloud-resolving model (CRM) simulation using open boundary conditions. The comparison of the SCM results with the reference CRM simulation provided insight into the ability of current convection and cloud schemes to represent organized convection. The CRM results enabled a detailed evaluation of the SCMs in terms of the thermodynamic structure and the convective mass flux of the system, the latter being closely related to the surface convective precipitation. It is shown that the SCMs could reproduce reasonably well the time evolution of the surface convective and stratiform precipitation, the convective mass flux, and the thermodynamic structure of the squall-line system. The thermodynamic structure simulated by the SCMs depended on how the models partitioned the precipitation between convective and stratiform. However, structural differences persisted in the thermodynamic profiles simulated by the SCMs and the CRM. These differences could be attributed to the fact that the total mass flux used to compute the SCM forcing differed from the convective mass flux. The SCMs could not adequately represent these organized mesoscale circulations and the microphysicallradiative forcing associated with the stratiform region. This issue is generally known as the ‘scale-interaction’ problem that can only be properly addressed in fully three-dimensional simulations. Sensitivity simulations run by several groups showed that the time evolution of the surface convective precipitation was considerably smoothed when the convective closure was based on convective available potential energy instead of moisture convergence. Finally, additional SCM simulations without using a convection parametrization indicated that the impact of a convection parametrization in forced SCM runs was more visible in the moisture profiles than in the temperature profiles because convective transport was particularly important in the moisture budget.

Evaluating forecasts from SETAR models of exchange rates

We consider the forecasting performance of two SETAR exchange rate models proposed by Kräger and Kugler [J. Int. Money Fin. 12 (1993) 195]. Assuming that the models are good approximations to the data generating process, we show that whether the non-linearities inherent in the data can be exploited to forecast better than a random walk depends on both how forecast accuracy is assessed and on the ‘state of nature’. Evaluation based on traditional measures, such as (root) mean squared forecast errors, may mask the superiority of the non-linear models. Generalized impulse response functions are also calculated as a means of portraying the asymmetric response to shocks implied by such models.

Testing the expectations theory of the term structure in threshold models

We test the expectations theory of the term structure of U.S. interest rates in nonlinear systems. These models allow the response of the change in short rates to past values of the spread to depend upon the level of the spread. The nonlinear system is tested against a linear system, and the results of testing the expectations theory in both models are contrasted. We find that the results of tests of the implications of the expectations theory depend on the size and sign of the spread. The long maturity spread predicts future changes of the short rate only when it is high.

Asymmetric output gap effects in Phillips Curve and mark-up pricing models: evidence for the US and the UK

A number of studies have found an asymmetric response of consumer price index inflation to the output gap in the US in simple Phillips curve models. We consider whether there are similar asymmetries in mark-up pricing models, that is, whether the mark-up over producers' costs also depends upon the sign of the (adjusted) output gap. The robustness of our findings to the price series is assessed, and also whether price-output responses in the UK are asymmetric.

Problems with binary pattern measures for flood model evaluation

As the calibration and evaluation of flood inundation models are a prerequisite for their successful application, there is a clear need to ensure that the performance measures that quantify how well models match the available observations are fit for purpose. This paper evaluates the binary pattern performance measures that are frequently used to compare flood inundation models with observations of flood extent. This evaluation considers whether these measures are able to calibrate and evaluate model predictions in a credible and consistent way, i.e. identifying the underlying model behaviour for a number of different purposes such as comparing models of floods of different magnitudes or on different catchments. Through theoretical examples, it is shown that the binary pattern measures are not consistent for floods of different sizes, such that for the same vertical error in water level, a model of a flood of large magnitude appears to perform better than a model of a smaller magnitude flood. Further, the commonly used Critical Success Index (usually referred to as F<2 >) is biased in favour of overprediction of the flood extent, and is also biased towards correctly predicting areas of the domain with smaller topographic gradients. Consequently, it is recommended that future studies consider carefully the implications of reporting conclusions using these performance measures. Additionally, future research should consider whether a more robust and consistent analysis could be achieved by using elevation comparison methods instead.

Mid-Holocene climates of the Americas: a dynamical response to changed seasonality

Simulations of the climatic response to mid-Holocene (6 ka BP) orbital forcing with two coupled ocean–atmosphere models (FOAM and CSM) show enhancement of monsoonal precipitation in parts of the American Southwest, Central America and northernmost South America during Northern Hemisphere summer. The enhanced onshore flow that brings precipitation into Central America is caused by a northward displacement of the inter-tropical convergence zone, driven by cooling of the equatorial and warming of the northern subtropical and mid-latitude ocean. Ocean feedbacks also enhance precipitation over the American Southwest, although the increase in monsoon precipitation there is largely driven by increases in land-surface temperature. The northward shift in the equatorial precipitation band that causes enhanced precipitation in Central America and the American Southwest has a negative feedback effect on monsoonal precipitation in northern South America. The simulations demonstrate that mid-Holocene aridity in the mid-continent of North America is dynamically linked to the orbitally induced enhancement of the summer monsoon in the American Southwest, with a spatial structure (wet in the Southwest and dry in the mid-continent) similar to that found in strong monsoon years today. Changes in winter precipitation along the west coast of North America, in Central America and along the Gulf Coast, caused by southward-displacement of the westerly storm tracks, indicate that changes in the Northern Hemisphere winter monsoon also play a role in regional climate changes during the mid-Holocene. Although the simulations with FOAM and CSM differ in detail, the general mechanisms and patterns are common to both. The model results thus provide a coherent dynamical explanation for regional patterns of increased or decreased aridity shown by vegetation, lake status and aeolian data from the Americas

Importance of initial state and atmospheric conditions for urban land surface models' performance

Urban land surface models (LSM) are commonly evaluated for short periods (a few weeks to months) because of limited observational data. This makes it difficult to distinguish the impact of initial conditions on model performance or to consider the response of a model to a range of possible atmospheric conditions. Drawing on results from the first urban LSM comparison, these two issues are considered. Assessment shows that the initial soil moisture has a substantial impact on the performance. Models initialised with soils that are too dry are not able to adjust their surface sensible and latent heat fluxes to realistic values until there is sufficient rainfall. Models initialised with too wet soils are not able to restrict their evaporation appropriately for periods in excess of a year. This has implications for short term evaluation studies and implies the need for soil moisture measurements to improve data assimilation and model initialisation. In contrast, initial conditions influencing the thermal storage have a much shorter adjustment timescale compared to soil moisture. Most models partition too much of the radiative energy at the surface into the sensible heat flux at the probable expense of the net storage heat flux.

Intercomparison of simulated global vegetation distribution in response to 6 kyr BP orbital forcing

The response of ten atmospheric general circulation models to orbital forcing at 6 kyr BP has been investigated using the BIOME model, which predicts equilibrium vegetation distribution, as a diagnostic. Several common features emerge: (a) reduced tropical rain forest as a consequence of increased aridity in the equatorial zone, (b) expansion of moisture-demanding vegetation in the Old World subtropics as a consequence of the expansion of the Afro–Asian monsoon, (c) an increase in warm grass/shrub in the Northern Hemisphere continental interiors in response to warming and enhanced aridity, and (d) a northward shift in the tundra–forest boundary in response to a warmer growing season at high northern latitudes. These broadscale features are consistent from model to model, but there are differences in their expression at a regional scale. Vegetation changes associated with monsoon enhancement and high-latitude summer warming are consistent with palaeoenvironmental observations, but the simulated shifts in vegetation belts are too small in both cases. Vegetation changes due to warmer and more arid conditions in the midcontinents of the Northern Hemisphere are consistent with palaeoenvironmental data from North America, but data from Eurasia suggests conditions were wetter at 6 kyr BP than today. The models show quantitatively similar vegetation changes in the intertropical zone, and in the northern and southern extratropics. The small differences among models in the magnitude of the global vegetation response are not related to differences in global or zonal climate averages, but reflect differences in simulated regional features. Regional-scale analyses will therefore be necessary to identify the underlying causes of such differences among models.

Impacts of 20th century aerosol emissions on the South Asian monsoon in the CMIP5 models

Comparison of single-forcing varieties of 20th century historical experiments in a subset of models from the Fifth Coupled Model Intercomparison Project (CMIP5) reveals that South Asian summer monsoon rainfall increases towards the present day in Greenhouse Gas (GHG)-only experiments with respect to pre-industrial levels, while it decreases in anthropogenic aerosol-only experiments. Comparison of these single-forcing experiments with the all-forcings historical experiment suggests aerosol emissions have dominated South Asian monsoon rainfall trends in recent decades, especially during the 1950s to 1970s. The variations in South Asian monsoon rainfall in these experiments follows approximately the time evolution of inter-hemispheric temperature gradient over the same period, suggesting a contribution from the large-scale background state relating to the asymmetric distribution of aerosol emissions about the equator. By examining the 24 available all-forcings historical experiments, we show that models including aerosol indirect effects dominate the negative rainfall trend. Indeed, models including only the direct radiative effect of aerosol show an increase in monsoon rainfall, consistent with the dominance of increasing greenhouse gas emissions and planetary warming on monsoon rainfall in those models. For South Asia, reduced rainfall in the models with indirect effects is related to decreased evaporation at the land surface rather than from anomalies in horizontal moisture flux, suggesting the impact of indirect effects on local aerosol emissions. This is confirmed by examination of aerosol loading and cloud droplet number trends over the South Asia region. Thus, while remote aerosols and their asymmetric distribution about the equator play a role in setting the inter-hemispheric temperature distribution on which the South Asian monsoon, as one of the global monsoons, operates, the addition of indirect aerosol effects acting on very local aerosol emissions also plays a role in declining monsoon rainfall. The disparity between the response of monsoon rainfall to increasing aerosol emissions in models containing direct aerosol effects only and those also containing indirect effects needs to be urgently investigated since the suggested future decline in Asian anthropogenic aerosol emissions inherent to the representative concentration pathways (RCPs) used for future climate projection may turn out to be optimistic. In addition, both groups of models show declining rainfall over China, also relating to local aerosol mechanisms. We hypothesize that aerosol emissions over China are large enough, in the CMIP5 models, to cause declining monsoon rainfall even in the absence of indirect aerosol effects. The same is not true for India.