Comparative connectomics reveals noncanonical wiring for color vision in human foveal retina.

The Old World macaque monkey and New World common marmoset provide fundamental models for human visual processing, yet the human ancestral lineage diverged from these monkey lineages over 25 Mya. We therefore asked whether fine-scale synaptic wiring in the nervous system is preserved across these three primate families, despite long periods of independent evolution. We applied connectomic electron microscopy to the specialized foveal retina where circuits for highest acuity and color vision reside. Synaptic motifs arising from the cone photoreceptor type sensitive to short (S) wavelengths and associated with "blue-yellow" (S-ON and S-OFF) color-coding circuitry were reconstructed. We found that distinctive circuitry arises from S cones for each of the three species. The S cones contacted neighboring L and M (long- and middle-wavelength sensitive) cones in humans, but such contacts were rare or absent in macaques and marmosets. We discovered a major S-OFF pathway in the human retina and established its absence in marmosets. Further, the S-ON and S-OFF chromatic pathways make excitatory-type synaptic contacts with L and M cone types in humans, but not in macaques or marmosets. Our results predict that early-stage chromatic signals are distinct in the human retina and imply that solving the human connectome at the nanoscale level of synaptic wiring will be critical for fully understanding the neural basis of human color vision.

Rare edges and abundant cores: range-wide variation in abundance in North American birds.

Understanding how the abundance of species varies across geographical ranges is central to ecology; however, few studies test hypotheses using detailed abundance estimates across the full ranges of species on a continental scale. Here, we use unprecedented, detailed estimates of breeding abundance for North American birds (eBird) to test two hypotheses for how abundance varies across species' ranges. We find widespread support for the rare-edge hypothesis-where the abundance of species declines near the range edge-reflecting both reduced occurrence and lower local abundance near range edges. By contrast, we find mixed support for the abundant-centre hypothesis-where the abundance of species peaks in the centre of the range and declines towards the edges-with limited support in conservative tests within species, but general support in among-species tests that control for unbalanced sampling and consider a broader definition of the range centre. Overall, results are consistent with a gradual decline in suitable conditions and increase in challenge towards the range edge that eventually limit the ability of populations to persist.

Is competitive ability the key adaptation to benign environments? Revisiting experiments on closely related species of tidal plants.

A central goal in biology is to understand which traits underlie adaptation to different environments. Yet, few studies have examined the relative contribution of competitive ability towards adaptive divergence among species occupying distinct environments. Here, we test the relative importance of competitive ability as an adaptation to relatively benign versus challenging environments, using previously published studies of closely related species pairs of primarily tidal plants subjected to reciprocal removal with transplant experiments in nature. Subordinate species typically occupy more challenging environments and showed consistent evidence for adaptation to challenging conditions, with no significant competitive effect on non-local, dominant species. In contrast, dominant species typically occupy relatively benign environments and performed significantly better than non-local, subordinate species that faced competition from the dominant species. Surprisingly, when the two species were not allowed to compete, the subordinate species performed as well as the dominant species in the benign environments where the subordinate species do not occur. These results suggest that competitive ability is the most important adaptation distinguishing the species that occupy relatively benign environments. The limited scope and number of suitable experimental studies encourage future work to test if these results are generalizable across taxa and environments.

Urban tolerance does not protect against population decline in North American birds.

Population declines of organisms are widespread and severe, but some species' populations have remained stable, or even increased. The reasons some species are less vulnerable to population decline than others are not well understood. Species that tolerate urban environments often have a broader environmental tolerance, which, along with their ability to tolerate one of the most human-modified habitats (i.e. cities), might allow them to persist in the face of diverse anthropogenic challenges. Here, we examined the relationship between urban tolerance and annual population trajectories for 397 North American bird species. Surprisingly, we found that urban tolerance was unrelated to species' population trajectories. The lack of a relationship between urban tolerance and population trajectories may reflect other factors driving population declines independent of urban tolerance, challenges that are amplified in cities (e.g. climate warming, disease), and other human impacts (e.g. conservation efforts, broad-scale land-use changes) that have benefitted some urban-avoidant species. Overall, our results illustrate that urban tolerance does not protect species against population decline.

Rapid adaptation of primate LGN neurons to drifting grating stimulation.

The visual system needs to dynamically adapt to changing environments. Much is known about the adaptive effects of constant stimulation over prolonged periods. However, there are open questions regarding adaptation to stimuli that are changing over time, interrupted, or repeated. Feature-specific adaptation to repeating stimuli has been shown to occur as early as primary visual cortex (V1), but there is also evidence for more generalized, fatigue-like adaptation that might occur at an earlier stage of processing. Here, we show adaptation in the lateral geniculate nucleus (LGN) of awake, fixating monkeys following brief (1 s) exposure to repeated cycles of a 4-Hz drifting grating. We examined the relative change of each neuron's response across successive (repeated) grating cycles. We found that neurons from all cell classes (parvocellular, magnocellular, and koniocellular) showed significant adaptation. However, only magnocellular neurons showed adaptation when responses were averaged to a population response. In contrast to firing rates, response variability was largely unaffected. Finally, adaptation was comparable between monocular and binocular stimulation, suggesting that rapid LGN adaptation is monocular in nature.NEW & NOTEWORTHY Neural adaptation can be defined as reduction of spiking responses following repeated or prolonged stimulation. Adaptation helps adjust neural responsiveness to avoid saturation and has been suggested to improve perceptual selectivity, information transmission, and predictive coding. Here, we report rapid adaptation to repeated cycles of gratings drifting over the receptive field of neurons at the earliest site of postretinal processing, the lateral geniculate nucleus of the thalamus.

Color as an Interspecific Badge of Status: A Comparative Test.

AbstractAnimals as diverse as cephalopods, insects, fish, and mammals signal social dominance to conspecifics to avoid costly fights. Even though between-species fights may be equally costly, the extent to which dominance signals are used between species is unknown. Here, we test the hypothesis that differences in color are associated with dominance between closely related species that aggressively interact over resources, examining between-species variation in colors that are used in within-species badges of status (black, white, and carotenoid coloration) in a comparative analysis of diverse species of birds. We found that dominant species have more black, on average, than subordinate species, particularly in regions important for aggressive signaling (face, throat, and bill). Furthermore, dominant species were more likely to have more black in comparisons in which the dominant species was similar in size or smaller than the subordinate, suggesting that black may be a more important signal when other signals of dominance (size) are missing. Carotenoid colors (i.e., red, pink, orange, and yellow) were not generally associated with dominance but may signal dominance in some taxonomic groups. White may have opposing functions: white was associated with dominance in species in which black was also associated with dominance but was associated with subordinance in species in which carotenoid-based dominance signals may be used. Overall, these results provide new evidence that colors may function broadly as signals of dominance among competing species. Such signals could help to mediate aggressive interactions among species, thereby reducing some costs of co-occurrence and facilitating coexistence in nature.

A Case for the "Competitive Exclusion-Tolerance Rule" as a General Cause of Species Turnover along Environmental Gradients.

AbstractClosely related, ecologically similar species often segregate their distributions along environmental gradients of time, space, and resources, but previous research suggests diverse underlying causes. Here, we review reciprocal removal studies in nature that experimentally test the role of interactions among species in determining their turnover along environmental gradients. We find consistent evidence for asymmetric exclusion coupled with differences in environmental tolerance causing the segregation of species pairs, where a dominant species excludes a subordinate from benign regions of the gradient but is unable to tolerate challenging regions to which the subordinate species is adapted. Subordinate species were consistently smaller and performed better in regions of the gradient typically occupied by the dominant species compared with their native distribution. These results extend previous ideas contrasting competitive ability with adaptation to abiotic stress to include a broader diversity of species interactions (intraguild predation, reproductive interference) and environmental gradients, including gradients of biotic challenge. Collectively, these findings suggest that adaptation to environmental challenge compromises performance in antagonistic interactions with ecologically similar species. The consistency of this pattern across diverse organisms, environments, and biomes suggests generalizable processes structuring the segregation of ecologically similar species along disparate environmental gradients, a phenomenon that we propose should be named the competitive exclusion-tolerance rule.

The Verriest Lecture: Pathways to color in the eye and brain.

In common with the majority of New World monkeys, marmosets show polymorphic color vision by allelic variation of X-chromosome genes encoding opsin pigments in the medium/long wavelength range. Male marmosets are thus obligate dichromats ("red-green color blind"), whereas females carrying distinct alleles on X chromosomes show one of three trichromatic phenotypes. Marmosets thus represent a "natural knock-out" system enabling comparison of red-green color vision in dichromatic and trichromatic visual systems. Further, study of short-wave (blue) cone pathways in marmosets has provided insights into primitive visual pathways for depth perception and attention. These investigations represent a parallel line to clinical research on color vision defects that was pioneered in studies by Guy Verreist, whom we honor in this eponymous lecture.

Among-species variation in hormone concentrations is associated with urban tolerance in birds.

As cities expand across the globe, understanding factors that underlie variation in urban tolerance is vital for predicting changes in patterns of biodiversity. Endocrine traits, like circulating hormone concentrations and regulation of endocrine responses, might contribute to variation in species' ability to cope with urban challenges. For example, variation in glucocorticoid and androgen concentrations has been linked to life-history and behavioural traits that are associated with urban tolerance. However, we lack an understanding of the degree to which evolved differences in endocrine traits predict variation in urban tolerance across species. We analysed 1391 estimates of circulating baseline corticosterone, stress-induced corticosterone, and testosterone concentrations paired with citizen-science-derived urban occurrence scores in a broad comparative analysis of endocrine phenotypes across 71 bird species that differ in their occurrence in urban habitats. Our results reveal context-dependent links between baseline corticosterone and urban tolerance, as well as testosterone and urban tolerance. Stress-induced corticosterone was not related to urban tolerance. These findings suggest that some endocrine phenotypes contribute to a species' tolerance of urban habitats, but also indicate that other aspects of the endocrine phenotype, such as the ability to appropriately attenuate responses to urban challenges, might be important for success in cities.

Experimental test of selection against hybridization as a driver of avian signal divergence.

Signal divergence may be pivotal in the generation and maintenance of new biodiversity by allowing closely related species to avoid some costs of co-occurrence. In birds, closely related, sympatric species are more divergent in their colour patterns than those that live apart, but the selective pressures driving this pattern remain unclear. Traditionally, signal divergence among sympatric species is thought to result from selection against hybridization, but broad evidence is lacking. Here, we conducted field experiments on naïve birds using spectrometer-matched, painted 3D-printed models to test whether selection against hybridization drives colour pattern divergence in the genus Poecile. To address selection for male colour pattern divergence without the influence of learning or the evolution of female discrimination in sympatry, we simulated secondary contact between Poecile species, and conducted mate choice experiments on naïve, allopatric females. We found that female black-capped chickadees (Poecile atricapillus) are equally likely to perform copulation solicitation displays to sympatric and allopatric heterospecific congeners when they are paired with conspecifics, but exhibit a strong preference for less divergent males when presented with paired heterospecific congeners. These results suggest that increased colour pattern divergence among sympatric species can reduce the likelihood of mixed mating in some contexts, and therefore should be favoured by selection against hybridization.

An observational study of self-reported migraine triggers and prospective evaluation of the relationships with occurrence of attacks enabled by a smartphone application (App).

OBJECTIVE: To investigate the relationship between self-reported triggers and the occurrence of migraine attacks using a smartphone application. BACKGROUND: One of several issues around the study of migraine attack triggers is that limited available evidence supports whether self-reported triggers can induce a headache on a particular subject. METHODS: This is an observational longitudinal cohort study of individuals with migraine registered to track their headaches prospectively using a smartphone application. For 90 days, participants entered daily data about triggers (potential triggers and premonitory symptoms) that may be associated with attack risk, as well as migraine symptoms. The statistical significance of univariate associations between each trigger and migraine recurrent events was determined for each individual. Statistically identified triggers were then compared to self-reported triggers. RESULTS: In 328 individuals (290/328 [88.4%] female; mean [standard deviation] 4.2 [1.5] migraine attacks/month) the mean (standard deviation) number of triggers moderately or highly endorsed per individual was 28.0 (7.7) in individuals presented with up to 38 possible triggers. Of these, an average (standard deviation) of 2.2 (2.1) triggers per individual were statistically associated with increased risk of attacks. Even the most commonly endorsed triggers (sleep quality, stress, tiredness/fatigue, sleep duration, dehydration, neck pain, missed meals, eyestrain, mean barometric pressure, and anxiety) were statistically associated in fewer than one third of individuals suspecting each, with the exception of neck pain (117/302 [38.7%]). CONCLUSIONS: Individuals with episodic migraine believe that many triggers contribute to their attacks; however, few of these withstand statistical testing at the individual level. Improved personal knowledge of potential triggers and premonitory symptoms may help individuals adopt behavioral changes to mitigate attack risk.

Analysis of Parvocellular and Magnocellular Visual Pathways in Human Retina.

Two main subcortical pathways serving conscious visual perception are the midget-parvocellular (P), and the parasol-magnocellular (M) pathways. It is generally accepted that the P pathway serves red-green color vision, but the relative contribution of P and M pathways to spatial vision is a long-standing and unresolved issue. Here, we mapped the spatial sampling properties of P and M pathways across the human retina. Data were obtained from immunolabeled vertical sections of six postmortem male and female human donor retinas and imaged using high-resolution microscopy. Cone photoreceptors, OFF-midget bipolar cells (P pathway), OFF-diffuse bipolar (DB) types DB3a and DB3b (M pathway), and ganglion cells were counted along the temporal horizontal meridian, taking foveal spatial distortions (postreceptoral displacements) into account. We found that the density of OFF-midget bipolar and OFF-midget ganglion cells can support one-to-one connections to 1.05-mm (3.6°) eccentricity. One-to-one connections of cones to OFF-midget bipolar cells are present to at least 10-mm (35°) eccentricity. The OFF-midget ganglion cell array acuity is well-matched to photopic spatial acuity measures throughout the central 35°, but the OFF-parasol array acuity is well below photopic spatial acuity, supporting the view that the P pathway underlies high-acuity spatial vision. Outside the fovea, array acuity of both OFF-midget and OFF-DB cells exceeds psychophysical measures of photopic spatial acuity. We conclude that parasol and midget pathway bipolar cells deliver high-acuity spatial signals to the inner plexiform layer, but outside the fovea, this spatial resolution is lost at the level of ganglion cells.SIGNIFICANCE STATEMENT We make accurate maps of the spatial density and distribution of neurons in the human retina to aid in understanding human spatial vision, interpretation of diagnostic tests, and the implementation of therapies for retinal diseases. Here, we map neurons involved with the midget-parvocellular (P pathway) and parasol-magnocellular (M pathway) through human retina. We find that P-type bipolar cells outnumber M-type bipolar cells at all eccentricities. We show that cone photoreceptors and P-type pathway bipolar cells are tightly connected throughout the retina, but that spatial resolution is lost at the level of the ganglion cells. Overall, the results support the view that the P pathway is specialized to serve both high acuity vision and red-green color vision.

Plasticity versus Evolutionary Divergence: What Causes Habitat Partitioning in Urban-Adapted Birds?

AbstractHabitat partitioning can facilitate the coexistence of closely related species and often results from competitive interference inducing plastic shifts of subordinate species in response to aggressive, dominant species (plasticity) or the evolution of ecological differences in subordinate species that reduce their ability to occupy habitats where the dominant species occurs (evolutionary divergence). Evidence consistent with both plasticity and evolutionary divergence exist, but the relative contributions of each to habitat partitioning have been difficult to discern. Here we use a global data set on the breeding occurrence of birds in cities to test predictions of these alternative hypotheses to explain previously described habitat partitioning associated with competitive interference. Consistent with plasticity, the presence of behaviorally dominant congeners in a city was associated with a 65% reduction in the occurrence of subordinate species, but only when the dominant was a widespread breeder in urban habitats. Consistent with evolutionary divergence, increased range-wide overlap with dominant congeners was associated with a 56% reduction in the occurrence of subordinates in cities, even when the dominant was absent from the city. Overall, our results suggest that both plasticity and evolutionary divergence play important, concurrent roles in habitat partitioning among closely related species in urban environments.

Experimental tests of selection against heterospecific aggression as a driver of avian colour pattern divergence.

Signal divergence is thought to reduce the costs of co-occurrence for closely related species and may thereby be important in the generation and maintenance of new biodiversity. In birds, closely related, sympatric species are more divergent in their colour patterns than those that live apart, but the selective pressures driving sympatric divergence in colour pattern are not well-understood. Here, we conducted field experiments on naïve birds using spectrometer-matched, painted, 3D-printed models to test whether selection against heterospecific aggression might drive colour pattern divergence in the genus Poecile. We found that territorial male black-capped chickadees (P. atricapillus) are equally likely to attack sympatric and allopatric congeners, and wintering flocks are equally likely to visit feeders occupied by sympatric and allopatric congeners, despite sympatric congeners being more divergent in colour pattern. These results suggest that either the concerted evolution of additional traits (e.g. discrimination), or interactions in sympatry that promote learning, is required if colour pattern divergence among sympatric species is to reduce heterospecific aggression. Alternatively, colour pattern divergence among sympatric species may be caused by other selective pressures, such as selection against hybridization or habitat partitioning and secondary signal adaptation.

Species interactions limit the occurrence of urban-adapted birds in cities.

Urbanization represents an extreme transformation of more natural systems. Populations of most species decline or disappear with urbanization, and yet some species persist and even thrive in cities. What determines which species persist or thrive in urban habitats? Direct competitive interactions among species can influence their distributions and resource use, particularly along gradients of environmental challenge. Given the challenges of urbanization, similar interactions may be important for determining which species persist or thrive in cities; however, their role remains poorly understood. Here, we use a global dataset to test among three alternative hypotheses for how direct competitive interactions and behavioral dominance may influence the breeding occurrence of birds in cities. We find evidence to support the competitive interference hypothesis: behaviorally dominant species were more widespread in urban habitats than closely related subordinate species, but only in taxa that thrive in urban environments (hereafter, urban adapted), and only when dominant and subordinate species overlapped their geographic ranges. This result was evident across diverse phylogenetic groups but varied significantly with a country's level of economic development. Urban-adapted, dominant species were more widespread than closely related subordinate species in cities in developed, but not developing, countries; countries in economic transition showed an intermediate pattern. Our results provide evidence that competitive interactions broadly influence species responses to urbanization, and that these interactions have asymmetric effects on subordinate species that otherwise could be widespread in urban environments. Results further suggest that economic development might accentuate the consequences of competitive interactions, thereby reducing local diversity in cities.

Fractal spike dynamics and neuronal coupling in the primate visual system.

KEY POINTS: We measured fractal (self-similar) fluctuations in ongoing spiking activity in subcortical (lateral geniculate nucleus, LGN) and cortical (area MT) visual areas in anaesthetised marmosets. Cells in the evolutionary ancient koniocellular LGN pathway and in area MT show high-amplitude fractal fluctuations, whereas evolutionarily newer parvocellular and magnocellular LGN cells do not. Spiking activity in koniocellular cells and MT cells shows substantial correlation to the local population activity, whereas activity in parvocellular and magnocellular cells is less correlated with local activity. We develop a model consisting of a fractal process and a global rate modulation which can reproduce and explain the fundamental relationship between fractal fluctuations and population coupling in LGN and MT. The model provides a unified account of apparently disparate aspects of neural spiking activity and can improve our understanding of information processing in evolutionary ancient and modern visual pathways. ABSTRACT: The brain represents and processes information through patterns of spiking activity, which is influenced by local and widescale brain circuits as well as intrinsic neural dynamics. Whether these influences have independent or linked effects on spiking activity is, however, not known. Here we measured spiking activity in two visual centres, the lateral geniculate nucleus (LGN) and cortical area MT, in marmoset monkeys. By combining the Fano-factor time curve, power spectral analysis and rescaled range analysis, we reveal inherent fractal fluctuations of spiking activity in LGN and MT. We found that the evolutionary ancient koniocellular (K) pathway in LGN and area MT exhibits strong fractal fluctuations at short (<1 s) time scales. Parvocellular (P) and magnocellular (M) LGN cells show weaker fractal fluctuations at longer (multi-second) time scales. In both LGN and MT, the amplitude and time scale of fractal fluctuations can explain short and long time scale spiking dynamics. We further show differential neuronal coupling of LGN and MT cells to local population spiking activity. The population coupling is intrinsically linked to fractal fluctuations: neurons showing stronger fluctuations are more strongly correlated to the local population activity. To understand this relationship, we modelled spiking activity using a fractal inhomogeneous Poisson process with dynamic rate, which is the product of an intrinsic stochastic fractal rate and a global modulatory gain. Our model explains the intrinsic links between neuronal spike rate and population coupling in LGN and MT, and establishes a unified account of dynamic spiking properties in afferent visual pathways.

Size-dependent costs of migration: Migrant bird species are subordinate to residents, but only at small body sizes.

Migrant species are commonly thought to be poor competitors in aggressive interactions with resident species. However, no studies have tested whether this relationship is widespread. Here, we compare the behavioural dominance of closely related species of migratory and nonmigratory birds, testing whether migrants are consistently subordinate to resident species in aggressive contests. We compiled published behavioural dominance data involving migrant and resident congeners, gathering additional data on the body mass and migratory distance of each species. Focal species included a diverse array of birds (28 taxonomic families, 12 orders) from around the world. We found that migrant species are usually subordinate to resident species, but that this relationship disappears at larger body sizes. For smaller birds (<500 g), resident species were behaviourally dominant in 83%-88% of comparisons; for larger birds (>500 g), resident species were dominant in only 25%-30% of comparisons. The relative difference in body mass best predicted dominance relationships among species, with larger species dominant in 80%-84% of comparisons. When migrant and resident masses were equal, however, resident species were still more likely to be dominant in smaller birds, suggesting that other factors may also contribute to the subordinate status of migrants. Overall, our results suggest that in smaller species, the evolution of migration is associated with lighter weights and other traits that compromise the competitive abilities of migrants relative to residents. In contrast, larger species appear able to evolve migration without compromising their size or competitive abilities in aggressive contests, suggesting size-dependent constraints on the evolution of migration.

Triggers of Primary Headaches: Issues and Pathways Forward.

The triggers of primary headaches have considerable significance for our understanding and management of headache and migraine. Triggers explain the variance in headaches - why they occur when they do. Trigger management is generally viewed as an important component of a comprehensive treatment approach for headaches. Historically, however, triggers have not had a prominent place in the headache literature. This situation began to change 20 to 30 years ago, and the pace of change has increased exponentially in recent times. Nevertheless, the field is beset with issues that have held it back from achieving more. This review will focus on elaborating those issues with the goal of suggesting ways forward. The first issue considered will be the definition of a trigger, and how specific triggers are labeled. Consideration will then be given to a classification system for triggers. The review will discuss next the evidence relating to whether self-reported triggers can, indeed, precipitate headaches, and how the capacity to elicit headaches may be acquired or extinguished. Attention will be given to the very important clinical issue of trigger management. Finally, the pathways forward will be proposed. Perhaps the most useful thing to accomplish at this point in time would be agreement on a definition of headache triggers, a list of triggers, and a classification system for triggers. This would greatly assist in comparing research on triggers from different research groups as well as eliminating some of the issues identified in this review. An authoritative body such as the American Headache Society or the International Headache Society, could establish a multidisciplinary committee that would complete these tasks. Consideration should also be given to incorporating triggers into the International Classification of Headache Disorders as an axis or via the use of codes, as this would raise the profile of triggers in assessment and management.

Types of Triggers in Migraine - Factor Structure of the Headache Triggers Sensitivity and Avoidance Questionnaire and Development of a New Short Form (HTSAQ-SF).

OBJECTIVE: To examine the factor structure of the Headache Triggers Sensitivity and Avoidance Questionnaire (HTSAQ) and its German version (HTSAQ-G), in order to identify potential different types of triggers. Furthermore, a short form of the questionnaire was developed. BACKGROUND: The HTSAQ includes 24 of the most commonly reported headache triggers (eg, stress, odors, lack of sleep). Both the HTSAQ and HTSAQ-G appeared to be reliable and valid measures of sensitivity to and avoidance of headache triggers. METHODS: In a cross-country collaboration, data from 2 cross-sectional studies including N = 391 individuals diagnosed with migraine from Australia (n = 222) and Germany (n = 169) were analyzed. The factor structure of the questionnaire was examined using exploratory and confirmatory factor analysis. Finally, a short form of the HTSAQ was constructed and evaluated regarding psychometric properties. RESULTS: Factor analytic results showed a differentiation between internal and external headache triggers, and different patterns of strategies in coping with triggers. The scales of both the original questionnaire as well as the developed short form showed good reliability (Cronbach's α = 0.76 to 0.96). As expected, negative correlations (r = -0.10 to -0.30, P = .006 to .044) with acceptance of pain were observed. Participants with chronic migraine showed significantly higher triggers sensitivity and avoidance of triggers than those with episodic migraine (t(389) = -9.12, P < .001, Cohens d = 0.93). CONCLUSIONS: Both the long and short forms of the questionnaire appear to be reliable and valid measures. The development of the short form of the questionnaire simplifies the use of the HTSAQ in clinical practice. Further research should focus on other primary headache disorders, such as tension-type headache or cluster headache.

Receptive Field Properties of Koniocellular On/Off Neurons in the Lateral Geniculate Nucleus of Marmoset Monkeys.

The koniocellular (K) layers of the primate dorsal lateral geniculate nucleus house a variety of visual receptive field types, not all of which have been fully characterized. Here we made single-cell recordings targeted to the K layers of diurnal New World monkeys (marmosets). A subset of recorded cells was excited by both increments and decrements of light intensity (on/off-cells). Histological reconstruction of the location of these cells confirmed that they are segregated to K layers; we therefore refer to these cells as K-on/off cells. The K-on/off cells show high contrast sensitivity, strong bandpass spatial frequency tuning, and their response magnitude is strongly reduced by stimuli larger than the excitatory receptive field (silent suppressive surrounds). Stationary counterphase gratings evoke unmodulated spike rate increases or frequency-doubled responses in K-on/off cells; such responses are largely independent of grating spatial phase. The K-on/off cells are not orientation or direction selective. Some (but not all) properties of K-on/off cells are consistent with those of local-edge-detector/impressed-by-contrast cells reported in studies of cat retina and geniculate, and broad-thorny ganglion cells recorded in macaque monkey retina. The receptive field properties of K-on/off cells and their preferential location in the ventral K layers (K1 and K2) make them good candidates for the direct projection from geniculate to extrastriate cortical area MT/V5. If so, they could contribute to visual information processing in the dorsal ("where" or "action") visual stream.SIGNIFICANCE STATEMENT We characterize cells in an evolutionary ancient part of the visual pathway in primates. The cells are located in the lateral geniculate nucleus (the main visual afferent relay nucleus), in regions called koniocellular layers that are known to project to extrastriate visual areas as well as primary visual cortex. The cells show high contrast sensitivity and rapid, transient responses to light onset and offset. Their properties suggest they could contribute to visual processing in the dorsal ("where" or "action") visual stream.