A newly recognized theropod assemblage from the Lewisville Formation (Woodbine Group; Cenomanian) and its implications for understanding Late Cretaceous Appalachian terrestrial ecosystems.

While the terrestrial fossil record of the mid-Cretaceous interval (Aptian to Cenomanian) in North America has been poorly studied, the recent focus on fossil localities from the western United States has offered a more detailed picture of vertebrate diversity, ecosystem dynamics and faunal turnover that took place on the western landmass of Laramidia. This is in stark contrast to the terrestrial record from the eastern landmass of Appalachia, where vertebrate fossils are rare and consist mostly of isolated and fragmentary remains. However, a detailed understanding of these fossil communities during this interval is necessary for comparison of the faunal patterns that developed during the opening of the Western Interior Seaway (WIS). The Woodbine Group of Texas is a Cenomanian age (95-100 mya) deposit consisting of shallow marine, deltaic, and terrestrial communities, which were only recently separated from their western counterparts. These deposits have yielded a wealth of vertebrate remains, yet non-avian theropods are still largely unknown. Recently, multiple localities in the Lewisville Formation of the Woodbine Group have yielded new non-avian theropod material, including numerous isolated teeth and postcranial remains. While largely fragmentary, this material is sufficiently diagnostic to identify the following taxa: a large-bodied carcharodontosaur, a mid-sized tyrannosauroid, a large ornithomimosaur, a large dromaeosaurine, a small dromaeosaurid, a small troodontid, and a small coelurosaur. Some of these groups represent the first occurrence for Appalachia and provide a broader understanding of a newly expanded faunal diversity for the Eastern landmass. The Lewisville Formation theropod fauna is similar in taxonomic composition to contemporaneous deposits in Laramidia, confirming that these groups were widespread across the continent prior to extension of the WIS. The Lewisville Formation documents the transitional nature of Cenomanian coastal ecosystems in Texas while providing additional details on the evolution of Appalachian communities shortly after WIS extension.

An early bothremydid from the Arlington Archosaur Site of Texas.

Four turtle taxa are previously documented from the Cenomanian Arlington Archosaur Site (AAS) of the Lewisville Formation (Woodbine Group) in Texas. Herein, we describe a new side-necked turtle (Pleurodira), Pleurochayah appalachius gen. et sp. nov., which is a basal member of the Bothremydidae. Pleurochayah appalachius gen. et sp. nov. shares synapomorphic characters with other bothremydids, including shared traits with Kurmademydini and Cearachelyini, but has a unique combination of skull and shell traits. The new taxon is significant because it is the oldest crown pleurodiran turtle from North America and Laurasia, predating bothremynines Algorachelus peregrinus and Paiutemys tibert from Europe and North America respectively. This discovery also documents the oldest evidence of dispersal of crown Pleurodira from Gondwana to Laurasia. Pleurochayah appalachius gen. et sp. nov. is compared to previously described fossil pleurodires, placed in a modified phylogenetic analysis of pelomedusoid turtles, and discussed in the context of pleurodiran distribution in the mid-Cretaceous. Its unique combination of characters demonstrates marine adaptation and dispersal capability among basal bothremydids.

An Enigmatic Small Neosuchian Crocodyliform from the Woodbine Formation of Texas.

New discoveries at the Arlington Archosaur Site (AAS), a Cenomanian (Late Cretaceous) locality in north-central Texas, are filling gaps in our knowledge of mid-Cretaceous Appalachian ecosystems, which remain poorly characterized. The AAS is notable because it preserves a diverse crocodyliform record. As seen in other sites that preserve four or more crocodyliform taxa, the species present at the AAS exhibit different snout shapes and body sizes, indicating that this high diversity of sympatric species was likely sustainable due to niche partitioning. Here we describe Scolomastax sahlsteini gen. et sp. nov., a new species of crocodyliform from the AAS, currently known from a partial right mandibular ramus. This species differs from other crocodyliforms in possessing features associated with durophagy or omnivory, including a shortened mandible, reduced tooth count, heterodonty, a dorsally expanded surangular, and enlarged attachments for jaw adductor muscles. Our phylogenetic analysis places this new taxon within Eusuchia as a member of Paralligatoridae and sister taxon to Paralligator gradilifrons. Scolomastax sahlsteini extends the record of paralligatorids into the Late Cretaceous of North America. This discovery represents the first appearance of this clade on the poorly known landmass of Appalachia, supporting a biogeographic connection between North America and Asia in the Early Cretaceous prior to completion of the Western Interior Seaway. However, relationships among other endemic crocodyliforms and tree instability within Paralligatoridae suggest further analysis is needed to resolve phylogenetic and biogeographic relationships (http://zoobank.org/urn:lsid:zoobank.org:pub:DC114471-6687-4BB5-8FAE-96F7278B1DAF). Anat Rec, 303:801-812, 2020. © 2019 Wiley Periodicals, Inc.

Quantifying shape and ecology in avian pedal claws: The relationship between the bony core and keratinous sheath.

Terrestrial tetrapods use their claws to interact with their environments in a plethora of ways. Birds in particular have developed a diversity of claw shapes since they are often not bound to terrestrial locomotion and have heterogeneous body masses ranging several orders of magnitude. Numerous previous studies have hypothesized a connection between pedal claw shape and ecological mode in birds, yet have generated conflicting results, spanning from clear ecological groupings based on claw shape to a complete overlap of ecological modes. The majority of these studies have relied on traditional morphometric arc measurements of keratinous sheaths and have variably accounted for likely confounding factors such as body mass and phylogenetic relatedness. To better address the hypothesized relationship between ecology and claw shape in birds, we collected 580 radiographs allowing visualization of the bony core and keratinous sheath shape in 21 avian orders. Geometric morphometrics was used to quantify bony core and keratinous sheath shape and was compared to results using traditional arc measurements. Neither approach significantly separates bird claws into coarse ecological categories after integrating body size and phylogenetic relatedness; however, some separation between ecological groups is evident and we find a gradual shift from the claw shape of ground-dwelling birds to those of predatory birds. Further, the bony claw core and keratinous sheath are significantly correlated, and the degree of functional integration does not differ across ecological groups. Therefore, it is likely possible to compare fossil bony cores with extant keratinous sheaths after applying corrections. Finally, traditional metrics and geometric morphometric shape are significantly, yet loosely correlated. Based on these results, future workers are encouraged to use geometric morphometric approaches to study claw geometry and account for confounding factors such as body size, phylogeny, and individual variation prior to predicting ecology in fossil taxa.

An unusual association of hadrosaur and therizinosaur tracks within Late Cretaceous rocks of Denali National Park, Alaska.

We report details of a unique association of hadrosaur and therizinosaur tracks found in the Late Cretaceous lower Cantwell Formation, Denali National Park, central Alaska Range, Alaska. This rock unit is now well-documented as a source of thousands of fossil footprints of vertebrates such as fishes, pterosaurs, and avialan and non-avialan dinosaurs. The lower Cantwell Formation in this area consists of numerous fining-upward successions of conglomerates and pebbly sandstones, cross-stratified and massive sandstones, interbedded sandstones and siltstones, organic-rich siltstones and shales, and rare, thin, bentonites, typically bounded by thin coal seams, and it contains a diverse fossil flora. We report the first North American co-occurrence of tracks attributable to hadrosaurs and therizinosaurs in the lower Cantwell Formation. Although previously un-reported in North America, this association of hadrosaur and therizinosaur tracks is more characteristic of the correlative Nemegt Formation in central Asia, perhaps suggesting that parameters defining the continental ecosystem of central Asia were also present in this part of Alaska during the Latest Cretaceous.

New data towards the development of a comprehensive taphonomic framework for the Late Jurassic Cleveland-Lloyd Dinosaur Quarry, Central Utah.

The Cleveland-Lloyd Dinosaur Quarry (CLDQ) is the densest deposit of Jurassic theropod dinosaurs discovered to date. Unlike typical Jurassic bone deposits, it is dominated by the presence of Allosaurus fragilis. Since excavation began in the 1920s, numerous hypotheses have been put forward to explain the taphonomy of CLDQ, including a predator trap, a drought assemblage, and a poison spring. In an effort to reconcile the various interpretations of the quarry and reach a consensus on the depositional history of CLDQ, new data is required to develop a robust taphonomic framework congruent with all available data. Here we present two new data sets that aid in the development of such a robust taphonomic framework for CLDQ. First, x-ray fluorescence of CLDQ sediments indicate elevated barite and sulfide minerals relative to other sediments from the Morrison Formation in the region, suggesting an ephemeral environment dominated by periods of hypereutrophic conditions during bone accumulation. Second, the degree of abrasion and hydraulic equivalency of small bone fragments dispersed throughout the matrix were analyzed from CLDQ. Results of these analyses suggest that bone fragments are autochthonous or parautochthonous and are derived from bones deposited in the assemblage rather than transported. The variability in abrasion exhibited by the fragments is most parsimoniously explained by local periodic re-working and re-deposition during seasonal fluctuations throughout the duration of the quarry assemblage. Collectively, these data support previous interpretations that the CLDQ represents an attritional assemblage in a poorly-drained overbank deposit where vertebrate remains were introduced post-mortem to an ephemeral pond during flood conditions. Furthermore, while the elevated heavy metals detected at the Cleveland-Lloyd Dinosaur Quarry are not likely the primary driver for the accumulation of carcasses, they are likely the result of multiple sources; some metals may be derived from post-depositional and diagenetic processes, and others are potentially produced from an abundance of decomposing vertebrate carcasses. These new data help to support the inferred depositional environment of the quarry as an ephemeral pond, and represent a significant step in understanding the taphonomy of the bonebed and Late Jurassic paleoecology in this region.

Risk stratification, perioperative and periprocedural management of the patient receiving anticoagulant therapy.

As a result of the aging US population and the subsequent increase in the prevalence of coronary disease and atrial fibrillation, therapeutic use of anticoagulants has increased. Perioperative and periprocedural management of anticoagulated patients has become routine for anesthesiologists, who frequently mediate communication between the prescribing physician and the surgeon and assess the risks of both thromboembolic complications and hemorrhage. Data from randomized clinical trials on perioperative management of antithrombotic therapy are lacking. Therefore, clinical judgment is typically needed regarding decisions to continue, discontinue, bridge, or resume anticoagulation and regarding the time points when these events should occur in the perioperative period. In this review, we will discuss the most commonly used anticoagulants used in outpatient settings and discuss their management in the perioperative period. Special considerations for regional anesthesia and interventional pain procedures will also be reviewed.

Fluvial transport potential of shed and root-bearing dinosaur teeth from the late Jurassic Morrison Formation.

Shed dinosaur teeth are commonly collected microvertebrate remains that have been used for interpretations of dinosaur feeding behaviors, paleoecology, and population studies. However, such interpretations may be biased by taphonomic processes such as fluvial sorting influenced by tooth shape: shed teeth, removed from the skull during life, and teeth possessing roots, removed from the skull after death. As such, teeth may behave differently in fluvial systems due to their differences in shape. In order to determine the influence of fluvial processes on the preservation and distribution of shed and root-bearing dinosaur teeth, the hydrodynamic behaviors of high-density urethane resin casts of shed and root-bearing Allosaurus and Camarasaurus teeth were experimentally tested for relative transport distances at increasing flow velocities in an artificial fluviatile environment. Results show that tooth cast specimens exhibited comparable patterns of transport at lower velocities, though the shed Camarasaurus teeth transported considerably farther in medium to higher flow velocities. Two-Way ANOVA tests indicate significant differences in the mean transport distances of tooth casts oriented perpendicular to flow (p < 0.05) with varying tooth morphologies and flow velocities. The differences exhibited in the transportability of shed and root-bearing teeth has important implications for taphonomic reconstructions, as well as future studies on dinosaur population dynamics, paleoecology, and feeding behaviors.

A pilot trial of intravenous pamidronate for chronic low back pain.

Intravenous (i.v.) bisphosphonates relieve pain in conditions such as Paget's disease of bone, metastatic bone disease, and multiple myeloma. Based on positive findings from a prior case series, we conducted a randomized placebo-controlled study to assess the analgesic effect of i.v. pamidronate in subjects with chronic low back pain (CLBP) and evidence of degenerative disease of the spine. Four groups of 11 subjects (7 active, 4 placebo) were enrolled at escalating dose levels of 30, 60, 90, and 180 mg pamidronate (the latter administered as two 90 mg infusions). Primary outcomes were safety and change from baseline in average daily pain scores, recorded at 1, 2, 3, and 6 months postinfusion using electronic diaries. Secondary outcomes included responder rate, daily worst pain, and pain-related interference with daily function. There were no pamidronate-related serious adverse events or other significant safety findings. A statistically significant overall treatment difference in pain scores was observed, with clinically meaningful effects persisting for 6 months in the 180 mg pamidronate group. Least squares mean changes in daily average pain score were -1.39 (SE=0.43) for placebo, and -1.53 (0.71), -1.26 (0.81), -1.42 (0.65), and -4.13 (0.65) for pamidronate 30, 60, 90, and 180 mg, respectively (P=0.012 for pamidronate 180 mg vs placebo). The proportion of responders, changes in worst pain, and pain interference with daily function were also significantly improved for pamidronate 180 mg compared with placebo. In conclusion, i.v. pamidronate, administered as two 90 mg infusions, decreased pain intensity for 6 months in subjects with CLBP.

Tectal etiology for irrepressible saccades: a case study in a Rhesus monkey.

Brain circuits controlling eye movements are widely distributed and complex. The etiology of irrepressible square wave saccades is not fully understood and is likely different for different neuropathologies. In a previous study, spontaneously occurring irrepressible saccades were noted after a cerebrovascular accident that damaged the rostral superior colliculus (SC) and its commissure in a Rhesus monkey. Here, we tracked and quantified the development of similar symptoms in a Rhesus monkey caused by a lesion in the rostromedial SC and its commissure. We documented the changes in these saccadic intrusions while the monkey attempted fixation of a target on three consecutive days post-onset. On the first day, eye jerk amplitude was ~10 degrees and the direction was ~30 degrees above the left horizontal meridian. On the second day, the amplitude decreased to 6.5 degrees and the direction shifted towards vertical, ~20 degrees to the left of the vertical meridian. Size, but not direction, of the eye jerks continued to decrease until intrusions dissipated within one month. Histological examination after ~6 months from the first appearance of the intrusions revealed a lesion in the commissure of the SC. Results from this and the previous study confirm the involvement of the commissure of the SC as the common target for triggering this neuropathy. Our data suggest that commissural fibers play an important role in maintaining normal visual stability. Interrupting the commissure between the two superior colliculi causes saccadic intrusions in the form of irrepressible jerking of the eyes, probably by disrupting inhibitory signals transmitted through the commissure. Furthermore, disappearance of the symptoms suggests that inhibitory fields within the SC are plastic and can expand, possibly via inputs from inter-collicular and nigrotectal pathways.

A flexible user-interface for audiovisual presentation and interactive control in neurobehavioral experiments.

A major problem facing behavioral neuroscientists is a lack of unified, vendor-distributed data acquisition systems that allow stimulus presentation and behavioral monitoring while recording neural activity. Numerous systems perform one of these tasks well independently, but to our knowledge, a useful package with a straightforward user interface does not exist. Here we describe the development of a flexible, script-based user interface that enables customization for real-time stimulus presentation, behavioral monitoring and data acquisition. The experimental design can also incorporate neural microstimulation paradigms. We used this interface to deliver multimodal, auditory and visual (images or video) stimuli to a nonhuman primate and acquire single-unit data. Our design is cost-effective and works well with commercially available hardware and software. Our design incorporates a script, providing high-level control of data acquisition via a sequencer running on a digital signal processor to enable behaviorally triggered control of the presentation of visual and auditory stimuli. Our experiments were conducted in combination with eye-tracking hardware. The script, however, is designed to be broadly useful to neuroscientists who may want to deliver stimuli of different modalities using any animal model.

Preventing post-thoracotomy pain syndrome.

This article provides a concise overview of post-thoracotomy pain syndrome, describes anesthetic and surgical factors that have been investigated to reduce the incidence of the syndrome, and explores the effectiveness of various treatments for this condition. Although some interventions (both procedural and pharmacologic) have been investigated in both preventing and treating post-thoracotomy pain syndrome, definitive studies are lacking and firm conclusions regarding the benefit of any intervention cannot be drawn. The problem is compounded further by our lack of understanding of the pathophysiologic mechanisms underlying the development of chronic pain after surgery. Going forward, it will be important to elucidate these mechanisms and conduct well-designed trials involving novel therapeutic agents for both prevention and treatment of post-thoracotomy pain syndrome.

Broad-scale patterns of late jurassic dinosaur paleoecology.

BACKGROUND: There have been numerous studies on dinosaur biogeographic distribution patterns. However, these distribution data have not yet been applied to ecological questions. Ecological studies of dinosaurs have tended to focus on reconstructing individual taxa, usually through comparisons to modern analogs. Fewer studies have sought to determine if the ecological structure of fossil assemblages is preserved and, if so, how dinosaur communities varied. Climate is a major component driving differences between communities. If the ecological structure of a fossil locality is preserved, we expect that dinosaur assemblages from similar environments will share a similar ecological structure. METHODOLOGY/PRINCIPAL FINDINGS: This study applies Ecological Structure Analysis (ESA) to a dataset of 100+ dinosaur taxa arranged into twelve composite fossil assemblages from around the world. Each assemblage was assigned a climate zone (biome) based on its location. Dinosaur taxa were placed into ecomorphological categories. The proportion of each category creates an ecological profile for the assemblage, which were compared using cluster and principal components analyses. Assemblages grouped according to biome, with most coming from arid or semi-arid/seasonal climates. Differences between assemblages are tied to the proportion of large high-browsing vs. small ground-foraging herbivores, which separates arid from semi-arid and moister environments, respectively. However, the effects of historical, taphonomic, and other environmental factors are still evident. CONCLUSIONS/SIGNIFICANCE: This study is the first to show that the general ecological structure of Late Jurassic dinosaur assemblages is preserved at large scales and can be assessed quantitatively. Despite a broad similarity of climatic conditions, a degree of ecological variation is observed between assemblages, from arid to moist. Taxonomic differences between Asia and the other regions demonstrate at least one case of ecosystem convergence. The proportion of different ecomorphs, which reflects the prevailing climatic and environmental conditions present during fossil deposition, may therefore be used to differentiate Late Jurassic dinosaur fossil assemblages. This method is broadly applicable to different taxa and times, allowing one to address questions of evolutionary, biogeographic, and climatic importance.

Current and emerging pharmacologic therapies for pain and challenges which still lay ahead.

This chapter seeks to provide a concise overview of the pharmacologic armamentarium available to treat pain. Drugs will be discussed in terms of their indications, mechanisms of action, and major side effects. For the purposes of this chapter, analgesics will be divided into two groups: current and emerging; current analgesics will be further subdivided into older analgesics and newer analgesics. Older analgesics will refer to drugs that have had FDA approval or were used off label for pain before 1990. Newer analgesics will refer to drugs developed or approved for treating pain since 1990. Finally, emerging analgesics will refer to drugs that have pre-clinical data or phase I/II data to suggest efficacy in treating pain but have not been validated by larger Phase III clinical trials. The chapter concludes with a chart that seeks to highlight current problems involved in pain pharmacotherapy.

Saccade trajectories evoked by sequential and colliding stimulation of the monkey superior colliculus.

Using microstimulation we employed an explicit experimental control of activity in the superior colliculus at two sites within the motor map. We compared saccade metrics and dynamics evoked at each site independently with those caused by sequential presentation and collisions of the two stimulation trains. Essentially, we forced controlled spatio-temporal patterns of activity into the saccade control circuit with various timing relationships from known sites within the collicular motor map, thus revealing the spatio-temporal transformation from superior colliculus to eye movement dynamics under experimentally controlled conditions. We extend prior findings about decreasing time intervals between sequential presentations of stimulations to include mid-flight combinations and dynamic modifications of trajectory. We explore how asynchronous collisions between two movements systematically engage a normalization mechanism of movement metrics, and demonstrate how dynamic patterns of activity across the SC motor map can create mid-flight curvature of movement through the post-collicular dynamics of a displacement controller. The explicit control addresses feasibility for systems control models and provides benchmark data for experimental verification of model mechanisms.

NGX-4010, a high-concentration capsaicin dermal patch for lasting relief of peripheral neuropathic pain.

NeurogesX Inc is developing NGX-4010, a rapid-delivery dermal patch application system that contains high-concentration trans-capsaicin, for the treatment of peripheral neuropathic pain. Capsaicin evokes a lasting and reversible refractory state in primary sensory neurons involved in the generation and maintenance of neuropathic pain. NGX-4010 can be applied to the painful skin area up to a total surface area of 1120 cm2. In phase I clinical trials, NGX-4010 increased the threshold for warmth detection, reduced epidermal sensory nerve fiber density and was well tolerated. In phase II trials, NGX-4010 was effective in reducing pain in patients with post-herpetic neuralgia (PHN), HIV-associated distal sensory neuropathy (HIV-DSP) and painful diabetic neuropathy (PDN). Data from phase III trials in patients with PHN demonstrated that significantly more pain relief was achieved by NGX-4010 (30 to 32% reduction from baseline) compared with a low-concentration capsaicin active control (20 to 24% reduction); however, only one of two studies involving patients with HIV-DSP met the primary endpoint. NGX-4010 appears to have the potential to be an effective adjunctive or a stand-alone therapy for PHN, as well as potentially for HIV-DSP and PDN. NGX-4010 has been granted approval by the European Commission and an NDA has been accepted for filing by the FDA.

Premotor inhibitory neurons carry signals related to saccade adaptation in the monkey.

Cerebellar output changes during motor learning. How these changes cause alterations of motoneuron activity and movement remains an unresolved question for voluntary movements. To answer this question, we examined premotor neurons for saccadic eye movement. Previous studies indicate that cells in the fastigial oculomotor region (FOR) within the cerebellar nuclei on one side exhibit a gradual increase in their saccade-related discharge as the amplitude of ipsiversive saccades adaptively decreases. This change in FOR activity could cause the adaptive change in saccade amplitude because neurons in the FOR project directly to the brain stem region containing premotor burst neurons (BNs). To test this possibility, we recorded the activity of saccade-related burst neurons in the area that houses premotor inhibitory burst neurons (IBNs) and examined their discharge during amplitude-reducing adaptation elicited by intrasaccadic target steps. We specifically analyzed their activity for off-direction (contraversive) saccades, in which the IBN activity would increase to reduce saccade size. Before adaptation, 29 of 42 BNs examined discharged, at least occasionally, for contraversive saccades. As the amplitude of contraversive saccades decreased adaptively, half of BNs with off-direction spike activity showed an increase in the number of spikes (14/29) or an earlier occurrence of spikes (7/14). BNs that were silent during off-direction saccades before adaptation remained silent after adaptation. These results indicate that the changes in the off-direction activity of BNs are closely related to adaptive changes in saccade size and are appropriate to cause these changes.

Differential hypertrophy and atrophy among all types of cutaneous innervation in the glabrous skin of the monkey hand during aging and naturally occurring type 2 diabetes.

Diabetic neuropathy (DN) is a common severe complication of type 2 diabetes. The symptoms of chronic pain, tingling, and numbness are generally attributed to small fiber dysfunction. However, little is known about the pathology among innervation to distal extremities, where symptoms start earliest and are most severe, and where the innervation density is the highest and includes a wide variety of large fiber sensory endings. Our study assessed the immunochemistry, morphology, and density of the nonvascular innervation in glabrous skin from the hands of aged nondiabetic rhesus monkeys and from age-matched monkeys that had different durations of spontaneously occurring type 2 diabetes. Age-related reductions occurred among all types of innervation, with epidermal C-fiber endings preferentially diminishing earlier than presumptive Adelta-fiber endings. In diabetic monkeys epidermal innervation density diminished faster, became more unevenly distributed, and lost immunodetectable expression of calcitonin gene-related peptide and capsaicin receptors, TrpV1. Pacinian corpuscles also deteriorated. However, during the first few years of hyperglycemia, a surprising hypertrophy occurred among terminal arbors of remaining epidermal endings. Hypertrophy also occurred among Meissner corpuscles and Merkel endings supplied by Abeta fibers. After longer-term hyperglycemia, Meissner corpuscle hypertrophy declined but the number of corpuscles remained higher than in age-matched nondiabetics. However, the diabetic Meissner corpuscles had an abnormal structure and immunochemistry. In contrast, the expanded Merkel innervation was reduced to age-matched nondiabetic levels. These results indicate that transient phases of substantial innervation remodeling occur during the progression of diabetes, with differential increases and decreases occurring among the varieties of innervation.

Distinct short-term and long-term adaptation to reduce saccade size in monkey.

In monkeys, saccades that repeatedly overshoot their targets adapt to become smaller by the time the monkey has made 1,000-2,000 saccades. In life, adaptation must keep movements accurate for long periods of time. Previous work describes only saccade adaptation that occurs within a few hours. Here we describe long-term saccade adaptation elicited in three monkeys by 19 days of training. Each day a monkey made saccades to track 16 degrees leftward and rightward target movements. During saccades, the target stepped back toward its starting position 6.4 degrees (40%) in two monkeys or 8 degrees (50%) in the third. After each day's adaptation, we blindfolded the monkey with goggles and returned it to its cage overnight. We found that adapting saccades for 19 days elicited significantly larger, long-lasting reduction in saccade size than did adapting for only 1 day. Further, after 19 days of adaptation we could elicit additional, apparently normal, short-term reduction in saccade size by increasing the size of the intra-saccade target movement. In contrast, we could elicit only small additional size reduction after only 1 day of adaptation. A simple model using separate short- and long-term adaptation mechanisms can reproduce many of the features of saccade gain exhibited by monkeys during a 19-day adaptation. We conclude that there is a long-term saccade-adaptation mechanism that is distinct from the well-characterized short-term system and that this newly recognized system is responsible for long-term maintenance of saccade accuracy.

Effect of visual error size on saccade adaptation in monkey.

Saccades that consistently over- or undershoot their targets gradually become smaller or larger, respectively. The signal that elicits adaptation of saccade size is a difference between eye and target positions appearing repeatedly at the ends of saccades. Here we describe how visual error size affects the size of saccade adaptation. At the end of each saccade, we imposed a constant-sized error by moving the target to a specified point relative to eye position. We tested a variety of error sizes imposed after saccades to target movements of 6, 12, and 18 degrees. We found that the size of the gain change elicited in a particular experiment depended on both the size of the imposed postsaccade error and on the size of the preceding target movement. For example, imposed errors of 4-5 degrees reduce saccades tracking 6, 12, and 18 degrees target movements by an average of 18, 35, and 45%, respectively. The most effective errors were those that were 15-45% of the size of the initial target eccentricity. Negative errors, which reduce saccade size, were more effective in changing saccade gain than were positive errors, which increased saccade size. For example, for 12 degrees target movements, negative and positive errors of 2-6 degrees changed saccade gain an average of 35 and 8%, respectively. This description of the relationship between error size and adaptation size improves our ability to adapt saccades in the laboratory and characterizes the error sizes that will best drive neurons carrying the adaptation-related visual error signal.