A common contrast pooling rule for suppression within and between the eyes

Recent work has revealed multiple pathways for cross-orientation suppression in cat and human vision. In particular, ipsiocular and interocular pathways appear to assert their influence before binocular summation in human but have different (1) spatial tuning, (2) temporal dependencies, and (3) adaptation after-effects. Here we use mask components that fall outside the excitatory passband of the detecting mechanism to investigate the rules for pooling multiple mask components within these pathways. We measured psychophysical contrast masking functions for vertical 1 cycle/deg sine-wave gratings in the presence of left or right oblique (645 deg) 3 cycles/deg mask gratings with contrast C%, or a plaid made from their sum, where each component (i) had contrast 0.5Ci%. Masks and targets were presented to two eyes (binocular), one eye (monoptic), or different eyes (dichoptic). Binocular-masking functions superimposed when plotted against C, but in the monoptic and dichoptic conditions, the grating produced slightly more suppression than the plaid when Ci $ 16%. We tested contrast gain control models involving two types of contrast combination on the denominator: (1) spatial pooling of the mask after a local nonlinearity (to calculate either root mean square contrast or energy) and (2) "linear suppression" (Holmes & Meese, 2004, Journal of Vision 4, 1080–1089), involving the linear sum of the mask component contrasts. Monoptic and dichoptic masking were typically better fit by the spatial pooling models, but binocular masking was not: it demanded strict linear summation of the Michelson contrast across mask orientation. Another scheme, in which suppressive pooling followed compressive contrast responses to the mask components (e.g., oriented cortical cells), was ruled out by all of our data. We conclude that the different processes that underlie monoptic and dichoptic masking use the same type of contrast pooling within their respective suppressive fields, but the effects do not sum to predict the binocular case.

Pooling and suppression in human spatial vision

A distinct feature of several recent models of contrast masking is that detecting mechanisms are divisively inhibited by a broadly tuned ‘gain pool’ of narrow-band spatial pattern mechanisms. The contrast gain control provided by this ‘cross-channel’ architecture achieves contrast normalisation of early pattern mechanisms, which is important for keeping them within the non-saturating part of their biological operating characteristic. These models superseded earlier ‘within-channel’ models, which had supposed that masking arose from direct stimulation of the detecting mechanism by the mask. To reveal the extent of masking, I measured the levels produced with large ranges of pattern spatial relationships that have not been explored before. Substantial interactions between channels tuned to different orientations and spatial frequencies were found. Differences in the masking levels produced with single and multiple component mask patterns provided insights into the summation rules within the gain pool. A widely used cross-channel masking model was tested on these data and was found to perform poorly. The model was developed and a version in which linear summation was allowed between all components within the gain pool but with the exception of the self-suppressing route typically provided the best account of the data. Subsequently, an adaptation paradigm was used to probe the processes underlying pooled responses in masking. This delivered less insight into the pooling than the other studies and areas were identified that require investigation for a new unifying model of masking and adaptation. In further experiments, levels of cross-channel masking were found to be greatly influenced by the spatio-temporal tuning of the channels involved. Old masking experiments and ideas relying on within-channel models were re-elevated in terms of contemporary cross-channel models (e.g. estimations of channel bandwidths from orientation masking functions) and this led to different conclusions than those originally arrived at. The investigation of effects with spatio-temporally superimposed patterns is focussed upon throughout this work, though it is shown how these enquiries might be extended to investigate effects across spatial and temporal position.

Measuring the spatial extent of texture pooling using reverse correlation

The local image representation produced by early stages of visual analysis is uninformative regarding spatially extensive textures and surfaces. We know little about the cortical algorithm used to combine local information over space, and still less about the area over which it can operate. But such operations are vital to support perception of real-world objects and scenes. Here, we deploy a novel reverse-correlation technique to measure the extent of spatial pooling for target regions of different areas placed either in the central visual field, or more peripherally. Stimuli were large arrays of micropatterns, with their contrasts perturbed individually on an interval-by-interval basis. By comparing trial-by-trial observer responses with the predictions of computational models, we show that substantial regions (up to 13 carrier cycles) of a stimulus can be monitored in parallel by summing contrast over area. This summing strategy is very different from the more widely assumed signal selection strategy (a MAX operation), and suggests that neural mechanisms representing extensive visual textures can be recruited by attention. We also demonstrate that template resolution is much less precise in the parafovea than in the fovea, consistent with recent accounts of crowding. © 2014 The Authors.

Do R&D strategies in high-tech sectors differ from those in low-tech sectors? An alternative approach to testing the pooling assumption

Most studies investigating the determinants of R&D investment consider pooled estimates. However, if the parameters are heterogeneous, pooled coefficients may not provide reliable estimates of individual industry effects. Hence pooled parameters may conceal valuable information that may help target government tools more efficiently across heterogeneous industries. There is little evidence to date on the decomposition of the determinants of R&D investment by industry. Moreover, the existing work does not distinguish between those R&D determinants for which pooling may be valid and those for which it is not. In this paper, we test the pooling assumption for a panel of manufacturing industries and find that pooling is valid only for output fluctuations, adjustment costs and interest rates. Implementing the test results into our model, we find government funding is significant only for low-tech R&D. Foreign R&D and skilled labour matter only in high-tech sectors. These results suggest important implications for R&D policy.

Service encounter conceptualisation:employees' service behaviours and customers' service quality perceptions

The enhancement of service quality is an area of optimal managerial relevance that has, to date, received minimal attention in the literature. Because customers' service quality evaluations are based almost entirely upon the behaviours of frontline employees, organisations rely heavily upon these employees to improve overall service quality provision. However, much of the literature looking at service quality enhancement lacks detail when examining the impact of employee service-related behaviours on customers' service quality perceptions. As a result, this paper comprehensively conceptualises those front-line employee behaviours which are the most likely to enhance customers' service quality perceptions. This conceptualisation is grounded in an extensive review of the services marketing literature, pooling together previously disparate research strands. Formal hypotheses are presented. Implications and future research directions are also discussed.

Macro determinants of FDI inflows to Japan:an analysis of source country characteristics

This paper examines the source country determinants of FDI into Japan. The paper highlights certain methodological and theoretical weaknesses in the previous literature and offers some explanations for hitherto ambiguous results. Specifically, the paper highlights the importance of panel data analysis, and the identification of fixed effects in the analysis rather than simply pooling the data. Indeed, we argue that many of the results reported elsewhere are a feature of this mis-specification. To this end, pooled, fixed effects and random effects estimates are compared. The results suggest that FDI into Japan is inversely related to trade flows, such that trade and FDI are substitutes. Moreover, the results also suggest that FDI increases with home country political and economic stability. The paper also shows that previously reported results, regarding the importance of exchange rates, relative borrowing costs and labour costs in explaining FDI flows, are sensitive to the econometric specification and estimation approach. The paper also discusses the importance of these results within a policy context. In recent years Japan has sought to attract FDI, though many firms still complain of barriers to inward investment penetration in Japan. The results show that cultural and geographic distance are only of marginal importance in explaining FDI, and that the results are consistent with the market-seeking explanation of FDI. As such, the attitude to risk in the source country is strongly related to the size of FDI flows to Japan. © 2007 The Authors Journal compilation © 2007 Blackwell Publishing Ltd.

Suppression and summation in contrast gain control for human vision

Over the last ten years our understanding of early spatial vision has improved enormously. The long-standing model of probability summation amongst multiple independent mechanisms with static output nonlinearities responsible for masking is obsolete. It has been replaced by a much more complex network of additive, suppressive, and facilitatory interactions and nonlinearities across eyes, area, spatial frequency, and orientation that extend well beyond the classical recep-tive field (CRF). A review of a substantial body of psychophysical work performed by ourselves (20 papers), and others, leads us to the following tentative account of the processing path for signal contrast. The first suppression stage is monocular, isotropic, non-adaptable, accelerates with RMS contrast, most potent for low spatial and high temporal frequencies, and extends slightly beyond the CRF. Second and third stages of suppression are difficult to disentangle but are possibly pre- and post-binocular summation, and involve components that are scale invariant, isotropic, anisotropic, chromatic, achromatic, adaptable, interocular, substantially larger than the CRF, and saturated by contrast. The monocular excitatory pathways begin with half-wave rectification, followed by a preliminary stage of half-binocular summation, a square-law transducer, full binocular summation, pooling over phase, cross-mechanism facilitatory interactions, additive noise, linear summation over area, and a slightly uncertain decision-maker. The purpose of each of these interactions is far from clear, but the system benefits from area and binocular summation of weak contrast signals as well as area and ocularity invariances above threshold (a herd of zebras doesn't change its contrast when it increases in number or when you close one eye). One of many remaining challenges is to determine the stage or stages of spatial tuning in the excitatory pathway.

The role of stochasticity in an information-optimal neural population code

In this paper we consider the optimisation of Shannon mutual information (MI) in the context of two model neural systems The first is a stochastic pooling network (population) of McCulloch-Pitts (MP) type neurons (logical threshold units) subject to stochastic forcing; the second is (in a rate coding paradigm) a population of neurons that each displays Poisson statistics (the so called 'Poisson neuron'). The mutual information is optimised as a function of a parameter that characterises the 'noise level'-in the MP array this parameter is the standard deviation of the noise, in the population of Poisson neurons it is the window length used to determine the spike count. In both systems we find that the emergent neural architecture and; hence, code that maximises the MI is strongly influenced by the noise level. Low noise levels leads to a heterogeneous distribution of neural parameters (diversity), whereas, medium to high noise levels result in the clustering of neural parameters into distinct groups that can be interpreted as subpopulations In both cases the number of subpopulations increases with a decrease in noise level. Our results suggest that subpopulations are a generic feature of an information optimal neural population.

Applying quantitative methods in the assessment of outcomes of pharmacotherapy of psoriasis

Healthcare providers and policy makers are faced with an ever-increasing number of medical publications. Searching for relevant information and keeping up to date with new research findings remains a constant challenge. It has been widely acknowledged that narrative reviews of the literature are susceptible to several types of bias and a systematic approach may protect against these biases. The aim of this thesis was to apply quantitative methods in the assessment of outcomes of topical therapies for psoriasis. In particular, to systematically examine the comparative efficacy, tolerability and cost-effectiveness of topical calcipotriol in the treatment of mild-to-moderate psoriasis. Over the years, a wide range of techniques have been used to evaluate the severity of psoriasis and the outcomes from treatment. This lack of standardisation complicates the direct comparison of results and ultimately the pooling of outcomes from different clinical trials. There is a clear requirement for more comprehensive tools for measuring drug efficacy and disease severity in psoriasis. Ideally, the outcome measures need to be simple, relevant, practical, and widely applicable, and the instruments should be reliable, valid and responsive. The results of the meta-analysis reported herein show that calcipotriol is an effective antipsoriatic agent. In the short-tenn, the pooled data found calcipotriol to be more effective than calcitriol, tacalcitol, coal tar and short-contact dithranol. Only potent corticosteroids appeared to have comparable efficacy, with less short-term side-effects. Potent corticosteroids also added to the antipsoriatic effect of calcipotriol, and appeared to suppress the occurrence of calcipotriol-induced irritation. There was insufficient evidence to support any large effects in favour of improvements in efficacy when calcipotriol is used in combination with systemic therapies in patients with severe psoriasis. However, there was a total absence of long-term morbidity data on the effectiveness of any of the interventions studied. Decision analysis showed that, from the perspective of the NHS as payer, the relatively small differences in efficacy between calcipotriol and short-contact dithranol lead to large differences in the direct cost of treating patients with mildto-moderate plaque psoriasis. Further research is needed to examine the clinical and economic issues affecting patients under treatment for psoriasis in the UK. In particular, the maintenance value and cost/benefit ratio for the various treatment strategies, and the assessment of patient's preferences has not yet been adequately addressed for this chronic recurring disease.

Contrast integration over area is extensive: a three-stage model of spatial summation

Classical studies of area summation measure contrast detection thresholds as a function of grating diameter. Unfortunately, (i) this approach is compromised by retinal inhomogeneity and (ii) it potentially confounds summation of signal with summation of internal noise. The Swiss cheese stimulus of T. S. Meese and R. J. Summers (2007) and the closely related Battenberg stimulus of T. S. Meese (2010) were designed to avoid these problems by keeping target diameter constant and modulating interdigitated checks of first-order carrier contrast within the stimulus region. This approach has revealed a contrast integration process with greater potency than the classical model of spatial probability summation. Here, we used Swiss cheese stimuli to investigate the spatial limits of contrast integration over a range of carrier frequencies (1–16 c/deg) and raised plaid modulator frequencies (0.25–32 cycles/check). Subthreshold summation for interdigitated carrier pairs remained strong (~4 to 6 dB) up to 4 to 8 cycles/check. Our computational analysis of these results implied linear signal combination (following square-law transduction) over either (i) 12 carrier cycles or more or (ii) 1.27 deg or more. Our model has three stages of summation: short-range summation within linear receptive fields, medium-range integration to compute contrast energy for multiple patches of the image, and long-range pooling of the contrast integrators by probability summation. Our analysis legitimizes the inclusion of widespread integration of signal (and noise) within hierarchical image processing models. It also confirms the individual differences in the spatial extent of integration that emerge from our approach.

Simulating the value of collaboration in multi-actor conservation planning

The loss of habitat and biodiversity worldwide has led to considerable resources being spent for conservation purposes on actions such as the acquisition and management of land, the rehabilitation of degraded habitats, and the purchase of easements from private landowners. Prioritising these actions is challenging due to the complexity of the problem and because there can be multiple actors undertaking conservation actions, often with divergent or partially overlapping objectives. We use a modelling framework to explore this issue with a study involving two agents sequentially purchasing land for conservation. We apply our model to simulated data using distributions taken from real data to simulate the cost of patches and the rarity and co-occurence of species. In our model each agent attempted to implement a conservation network that met its target for the minimum cost using the conservation planning software Marxan. We examine three scenarios where the conservation targets of the agents differ. The first scenario (called NGO-NGO) models the situation where two NGOs are both are targeting different sets of threatened species. The second and third scenarios (called NGO-Gov and Gov-NGO, respectively) represent a case where a government agency attempts to implement a complementary conservation network representing all species, while an NGO is focused on achieving additional protection for the most endangered species. For each of these scenarios we examined three types of interactions between agents: i) acting in isolation where the agents are attempting to achieve their targets solely though their own actions ii) sharing information where each agent is aware of the species representation achieved within the other agent’s conservation network and, iii) pooling resources where agents combine their resources and undertake conservation actions as a single entity. The latter two interactions represent different types of collaborations and in each scenario we determine the cost savings from sharing information or pooling resources. In each case we examined the utility of these interactions from the viewpoint of the combined conservation network resulting from both agents' actions, as well as from each agent’s individual perspective. The costs for each agent to achieve their objectives varied depending on the order in which the agents acted, the type of interaction between agents, and the specific goals of each agent. There were significant cost savings from increased collaboration via sharing information in the NGO-NGO scenario were the agent’s representation goals were mutually exclusive (in terms of specie targeted). In the NGO-Gov and Gov-NGO scenarios, collaboration generated much smaller savings. If the two agents collaborate by pooling resources there are multiple ways the total cost could be shared between both agents. For each scenario we investigate the costs and benefits for all possible cost sharing proportions. We find that there are a range of cost sharing proportions where both agents can benefit in the NGO-NGO scenarios while the NGO-Gov and Gov-NGO scenarios again showed little benefit. Although the model presented here has a range of simplifying assumptions, it demonstrates that the value of collaboration can vary significantly in different situations. In most cases, collaborating would have associated costs and these costs need to be weighed against the potential benefits from collaboration. The model demonstrates a method for determining the range of collaboration costs that would result in collaboration providing an efficient use of scarce conservation resources.

Spatially extensive summation of contrast-energy is revealed by contrast detection of micro-pattern textures

Vision must analyze the retinal image over both small and large areas to represent fine-scale spatial details and extensive textures. The long-range neuronal convergence that this implies might lead us to expect that contrast sensitivity should improve markedly with the contrast area of the image. But this is at odds with the orthodox view that contrast sensitivity is determined merely by probability summation over local independent detectors. To address this puzzle, I aimed to assess the summation of luminance contrast without the confounding influence of area-dependent internal noise. I measured contrast detection thresholds for novel Battenberg stimuli that had identical overall dimensions (to clamp the aggregation of noise) but were constructed from either dense or sparse arrays of micro-patterns. The results unveiled a three-stage visual hierarchy of contrast summation involving (i) spatial filtering, (ii) long-range summation of coherent textures, and (iii) pooling across orthogonal textures. Linear summation over local energy detectors was spatially extensive (as much as 16 cycles) at Stage 2, but the resulting model is also consistent with earlier classical results of contrast summation (J. G. Robson & N. Graham, 1981), where co-aggregation of internal noise has obscured these long-range interactions.

Orientation masking and cross-orientation suppression (XOS):implications for estimates of channel bandwidth

Most contemporary models of spatial vision include a cross-oriented route to suppression (masking from a broadly tuned inhibitory pool), which is most potent at low spatial and high temporal frequencies (T. S. Meese & D. J. Holmes, 2007). The influence of this pathway can elevate orientation-masking functions without exciting the target mechanism, and because early psychophysical estimates of filter bandwidth did not accommodate this, it is likely that they have been overestimated for this corner of stimulus space. Here we show that a transient 40% contrast mask causes substantial binocular threshold elevation for a transient vertical target, and this declines from a mask orientation of 0° to about 40° (indicating tuning), and then more gently to 90°, where it remains at a factor of ∼4. We also confirm that cross-orientation masking is diminished or abolished at high spatial frequencies and for sustained temporal modulation. We fitted a simple model of pedestal masking and cross-orientation suppression (XOS) to our data and those of G. C. Phillips and H. R. Wilson (1984) and found the dependency of orientation bandwidth on spatial frequency to be much less than previously supposed. An extension of our linear spatial pooling model of contrast gain control and dilution masking (T. S. Meese & R. J. Summers, 2007) is also shown to be consistent with our results using filter bandwidths of ±20°. Both models include tightly and broadly tuned components of divisive suppression. More generally, because XOS and/or dilution masking can affect the shape of orientation-masking curves, we caution that variations in bandwidth estimates might reflect variations in processes that have nothing to do with filter bandwidth.

Theory and data for area summation of contrast with and without uncertainty:evidence for a noisy energy model

Contrast sensitivity improves with the area of a sine-wave grating, but why? Here we assess this phenomenon against contemporary models involving spatial summation, probability summation, uncertainty, and stochastic noise. Using a two-interval forced-choice procedure we measured contrast sensitivity for circular patches of sine-wave gratings with various diameters that were blocked or interleaved across trials to produce low and high extrinsic uncertainty, respectively. Summation curves were steep initially, becoming shallower thereafter. For the smaller stimuli, sensitivity was slightly worse for the interleaved design than for the blocked design. Neither area nor blocking affected the slope of the psychometric function. We derived model predictions for noisy mechanisms and extrinsic uncertainty that was either low or high. The contrast transducer was either linear (c1.0) or nonlinear (c2.0), and pooling was either linear or a MAX operation. There was either no intrinsic uncertainty, or it was fixed or proportional to stimulus size. Of these 10 canonical models, only the nonlinear transducer with linear pooling (the noisy energy model) described the main forms of the data for both experimental designs. We also show how a cross-correlator can be modified to fit our results and provide a contemporary presentation of the relation between summation and the slope of the psychometric function.

Simulating the value of collaboration in multi-actor conservation planning

The loss of habitat and biodiversity worldwide has led to considerable resources being spent on conservation interventions. Prioritising these actions is challenging due to the complexity of the problem and because there can be multiple actors undertaking conservation actions, often with divergent or partially overlapping objectives. We explore this issue with a simulation study involving two agents sequentially purchasing land for the conservation of multiple species using three scenarios comprising either divergent or partially overlapping objectives between the agents. The first scenario investigates the situation where both agents are targeting different sets of threatened species. The second and third scenarios represent a case where a government agency attempts to implement a complementary conservation network representing 200 species, while a non-government organisation is focused on achieving additional protection for the ten rarest species. Simulated input data was generated using distributions taken from real data to model the cost of parcels, and the rarity and co-occurrence of species. We investigated three types of collaborative interactions between agents: acting in isolation, sharing information and pooling resources with the third option resulting in the agents combining their resources and effectively acting as a single entity. In each scenario we determine the cost savings when an agent moves from acting in isolation to either sharing information or pooling resources with the other agent. The model demonstrates how the value of collaboration can vary significantly in different situations. In most cases, collaborating would have associated costs and these costs need to be weighed against the potential benefits from collaboration. Our model demonstrates a method for determining the range of costs that would result in collaboration providing an efficient use of scarce conservation resources.