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Therapeutic product candidates
Factor VIII Intolerance
Haemophilia A (HA) is a rare chronic bleeding disorder which leads to inadequate clotting of the blood in response to any type of injury or surgery. The most common form of HA is congenital HA, which mostly affects males due to its X-linked recessive inheritance pattern and has an incidence of 1 in 5000.
There are approximately 33,000 HA patients under active management in the USA and Europe. The mainstay of HA treatment is the life-long replacement therapy with synthetic FVIII to achieve haemostasis using either plasma derived or recombinant FVIII products.
The most challenging complication of therapy is the development of anti-FVIII neutralising antibodies which occurs in 20-30% of patients. These antibodies block (inhibit) the pro-coagulant function of FVIII and are therefore termed inhibitors. The development of such inhibitors sharply decreases the efficacy of the FVIII and quickly renders the FVIII replacement ineffective and can result in joint damage, brain damage and, ultimately, death. There are currently no treatments available to prevent or treat the development of neutralising antibodies.
The ATX-F8-117 discovery programme has identified two Apitopes® which have the potential to treat and prevent Factor VIII inhibitor formation by as much as 96%.


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No Amendments Surface in House Finance Budget Hearing
The House Finance, Ways, and Means Committee took up UT's proposed budget on Monday, March 6. The Committee's tone was positive, although some questions did arise about recent comments by UT Knoxville Chancellor Beverly Davenport regarding the future of UTK's Office of Diversity and Inclusion. Committee members acknowledged and voiced support for UT's effort to … Continued
Bill Expanding Firearms On Campus Moves Forward
A measure expanding access to firearms on public college campuses passed the House Civil Justice Committee this week on a 6-3 party line vote. Sponsored by Rep. Dennis Powers (R-Jacksboro), House Bill 745 would permit part-time employees who have handgun carry permits to carry a concealed weapon on campus. The bill, which was originally narrowly … Continued
Senate Education Committee to Conduct UT Budget Hearing on Wednesday
The Senate Education Committee will conduct UT's budget hearing on Wednesday, March 1. In recent years, UT's budget hearings before this Senate Committee have been less focused on the budget proposal itself and more focused on issues facing the University that the Committee has deemed controversial. UT will concentrate budget testimony on efforts underway to … Continued
House Civil Justice Subcommittee to Consider Firearms Legislation with Broad Impact on Public Universities
As an increasing number of bills come before the House Civil Justice Subcommittee, two firearms bills, in particular, have a direct and broad impact on public colleges and universities in Tennessee. Both bills would have the effect of expanding the possession of firearms on public campuses to students. Public colleges and universities in Tennessee are not … Continued
UT Day on the Hill Showcases National Security Efforts
From working on nuclear security to operating a regional biocontainment laboratory to active shooter response training and more, the University of Tennessee is helping keep America safe. Every UT campus and institute plays a role in ensuring America's safety was the message students and faculty shared during today's 11th UT Day on the Hill. The … Continued


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We examined the motor error hypothesis of visual and auditory interaction in the superior colliculus (SC), first tested by Jay and Sparks in the monkey. We trained cats to direct their eyes to the location of acoustic sources and studied the effects of eye position on both the ability of cats to localize sounds and the auditory responses of SC neurons with the head restrained. Sound localization accuracy was generally not affected by initial eye position, i.e., accuracy was not proportionally affected by the deviation of the eyes from the primary position at the time of stimulus presentation, showing that eye position is taken into account when orienting to acoustic targets. The responses of most single SC neurons to acoustic stimuli in the intact cat were modulated by eye position in the direction consistent with the predictions of the "motor error" hypothesis, but the shift accounted for only two-thirds of the initial deviation of the eyes. However, when the average horizontal sound localization error, which was ∼35% of the target amplitude, was taken into account, the magnitude of the horizontal shifts in the SC auditory receptive fields matched the observed behavior. The modulation by eye position was not due to concomitant movements of the external ears, as confirmed by recordings carried out after immobilizing the pinnae of one cat. However, the pattern of modulation after pinnae immobilization was inconsistent with the observations in the intact cat, suggesting that, in the intact animal, information about the position of the pinnae may be taken into account.
In anesthetized animals, the various sensory and motor maps in the superior colliculus (SC) are in approximate alignment (Gordon 1973; King and Palmer 1983; Meredith and Stein 1985; Middlebrooks and Knudsen 1984; Stein and Clamann 1981; Stein and Meredith 1993), meaning that acoustic and visual stimuli at the same spatial locations stimulate the same part of the SC. The alignment of auditory and visual topography is thought to provide a substrate for sensorimotor integration (Harris et al. 1980; Stein and Meredith 1993). However, as first pointed out by Pöppel (1973), since the coordinate systems for vision and audition are different, with the visual map oculocentric and the auditory map head-centered, this alignment holds only when the two coordinate systems are aligned, i.e., when the eyes and head point in the same direction.
Thus in behaving animals, whose eyes are free to move in the head, one would expect the SC maps to become misaligned whenever the animal is not looking straight ahead. If this alignment is critical for sound localization or for bimodal integration, the consequence of misalignment between these two maps might be mislocalization of the source of a sound whenever the eyes are away from the primary position. Furthermore, the magnitude of the expected error in sound localization should be proportional to the deviation of the eyes from the primary position (Harris et al. 1980). This hypothesis can be tested behaviorally.
Alternatively, the collicular representation of auditory space could shift to compensate for changes in eye position to maintain the alignment of the receptive fields (RFs) across the different modalities. Such a shift would require a modulating signal proportional to the position of the eyes within the orbit to yield a representation of the eye motor error, which is the difference between eye position and the target. In this scheme, also known as the "motor error hypothesis" (Sparks 1986), the shifted auditory RFs would not encode the location of auditory sources relative to the head, but rather their location relative to the position of the eyes within the orbit. This hypothesis can be tested physiologically.
Jay and Sparks (1987) found evidence supporting the motor error hypothesis in the SC of the monkey: they found that the auditory response was modulated by eye position, although it only partly compensated for the motor error. More recently, Groh et al. (2001) and Zwiers et al. (2004) both found evidence of modulation by eye position in auditory responses by a variable number of cells in the inferior colliculus (IC) even though their monkeys were not trained to orient to the acoustic stimuli. However, in the cat, conflicting evidence has been presented. Harris et al. (1980) claim that cats do not move their eyes enough to require compensation, while Hartline et al. (1995) and Peck et al. (1995) provide evidence for compensation. Furthermore, shifts of auditory receptive fields in the SC of cats could result from changes in the acoustic input to the eardrums brought about by the highly mobile pinnae (Middlebrooks and Knudsen 1987). However, the interpretation of the available data in all three studies (Harris et al. 1980; Hartline et al. 1995; Peck et al. 1995) is hindered by limited sample size, untrained and inconsistent subjects, and inadequate monitoring of eye or ear position. We have designed this study to overcome these problems using cats who consistently localized sound by looking at the acoustic sources and well-controlled experimental conditions.
Our results show that most cells driven by acoustic stimuli in the deep and intermediate layers of the SC of the cat are modulated by eye position in a manner consonant with the motor error hypothesis. Although the shift in auditory RFs did not compensate fully for the motor error, it did match the behavioral motor error, which was calculated on the basis of the propensity for the saccades to auditory targets to undershoot the target. Finally, by recording from cells in the SC in a cat whose pinnae had been immobilized, we showed for the first time that the modulation cannot be due to the changes in acoustics brought about by movements of the pinnae of the cat while fixating different positions. However, the modulation seen in the pinna immobilized cat was not consistent with the modulation seen in the intact cat, suggesting that eye position, pinna position, and a motor efferent copy command may all play a role in the acoustic modulation in the SC. A preliminary account of portions of this work has been presented (Populin and Yin 1998a).
Subjects, surgery, and experimental setup
A detailed description of the materials, methods, and animal training procedures has been presented previously (Populin and Yin 1998b). Briefly, 10 cats were trained to localize sound by looking at the perceived location of the sources. Single neuronal recordings made in the SC of six of them are included in this report. Under sterile conditions, a recording cylinder was implanted on the midline of the skull to access the SC on both sides with vertically oriented electrode penetrations. The animals had already undergone a surgical procedure to implant eye (Judge et al. 1980) and pinna (Populin and Yin 1998c) coils and a post to restrain the head and had been trained on the behavioral tasks.
Anesthesia was first induced with an intramuscular injection of ketamine (20 mg/kg) and acepromazine (0.2 mg/kg), followed by intravenous sodium pentobarbital or gas (halothane or isoflurane). All surgical and experimental procedures employed were approved by the University of Wisconsin Animal Care Committee and were in accordance with the National Institutes of Health Guide for the Care and Use of Laboratory Animals.
The experiments were carried out in a dimly illuminated (2.2 × 2.5 × 2.5 m) single-walled sound-attenuated chamber with the inner walls and major pieces of equipment in it covered with 10.2-cm Sonex foam (Ilbruck, Minneapolis, MN) to attenuate acoustic reflections. All experiments were carried out under head-restrained conditions.
Eye and pinna movement recordings and stimuli presentation
Eye and pinna movements were recorded with the scleral search coil technique (Robinson 1963). The analog output of the CNC Engineering (Seattle, WA) system used to measure eye and pinna movements was digitally sampled at 500 Hz with an A/D converter and stored on disk for off-line analysis.
Acoustic stimuli were generated with a Digital Stimulus System (Rhode 1976) and presented via Radio Shack Super tweeters (model 40-1310A) that had been modified to transduce low frequencies. Visual stimuli were presented from small (2.0 mm diam) red (λ max = 635 nm) light-emitting diodes (LEDs) that subtended a visual angle of 0.2° and were mounted at the center of each speaker. The transducers were placed in the frontal hemifield, 62 cm away from the cat's head behind a dark curtain that allowed sounds to be heard and LEDs to be seen when lit. The standard acoustic stimulus was a 163-ms noise burst (0.1–25 kHz) with 7-ms rise-fall windows and was repeated as needed to produce bursts of longer durations. All aspects of data acquisition and stimulus presentation were controlled by a MicroVAX II minicomputer (Digital Equipment, Maynard, MA) running custom software.
Physiological recordings
Recording sessions started 5–7 days after the implantation of the recording cylinder, took place five to six times per week, and lasted about 3 h each. The animals were continuously monitored through closed circuit TV for signs of distress. The end of a recording session was typically signaled by the animal falling asleep or stopping due to satiety.
Parylene-covered tungsten microelectrodes (MicroProbe, Potomac, MD) were driven through the intact dura mater with an eccentric Narishige (Narishige International, East Meadow, NY) microdrive. Extracellular signals were amplified and band-pass filtered (300–3,000 Hz). A window discriminator (Bak Electronics, Mount Airy, MD) was used to generate pulses every time an action potential met the spatial and temporal criteria set by the investigator. These were recorded with 1-μs resolution with a custom-made event timer. All data presented in this report are from well-isolated single neurons.
Behavioral tasks
We initially trained the cats on a variety of visual and auditory oculomotor tasks (visual and auditory fixations, standard and delayed saccades, and sensory probes, as described in Populin and Yin 1998b) delivered in random order so that on any given trial the cat could not predict what the task would be. This training period took an average of 7 mo, at the end of which we did a sterile surgery and mounted a recording cylinder on the cat to commence physiological recordings from the SC.
The behavioral experiments presented below used the auditory standard saccade task (Fig. 1A). This task used a visual fixation stimulus (LED), which allowed us to control the initial position of the eyes, and an acoustic stimulus (SPEAKER), which was the target for the saccade. The cat was required to first acquire and maintain fixation on the fixation LED until it was turned off at the same time that the speaker was turned on and then make a saccade to the newly presented target and maintain fixation until the sound was turned off. A reward was delivered if temporal and spatial criteria were met. The rationale for establishing the spatial criteria for success is discussed in Populin and Yin (1998b).
During physiological recordings, to determine if the responses of SC auditory neurons were modulated by fixation position, we used the auditory sensory probe task (Fig. 1B). This task is a variation of the auditory delayed saccade task (Populin and Yin 1998b), which was used for training because it required the cats to treat acoustic stimuli as targets for saccadic eye movements. In the sensory probe task, the cat is required to fixate for a predetermined period of time on one of three to five fixation LEDs located along the horizontal axis. During this period of fixation, a broadband noise burst (probe SPEAKER), presented from another speaker, was used to probe the neuron's responses to the sound. To receive a food reward, the cat had to maintain its eyes within an electronic window, typically (≤6 × 6°), centered about the fixation LED until the end of the trial when both the fixation LED and probe speaker were turned off. Usually we varied one or more parameters (e.g., the fixation LED or the acoustic speaker location) during any particular recording session so that a variety of tasks was always presented and the cat could never anticipate what the upcoming trial would be.
For the auditory delayed saccade task (data not shown), the fixation led was not turned off until the auditory target had been on for 300–500 ms. The offset of the fixation LED signaled the cat to make a saccadic eye movement to the auditory target. It is important to note that the sensory probe and delayed saccade tasks are identical up to the time that the fixation LED is extinguished to signal the saccade. Ideally, we would have preferred to use the delayed saccade task for our recordings, but the best response area of most units was outside the oculomotor range of the cat, which is approximately ±20°. We are confident that the cat considered the probe speakers as potential saccade targets because when we switched to runs with auditory delayed saccades or auditory fixation tasks, they immediately made short-latency saccades to those targets.
Pinnae immobilization
Our behavioral studies showed that cats move their external ears in a systematic fashion in response to acoustic stimuli and when fixating at different spatial positions (Populin and Yin 1998c). Therefore the position of the ears varies as the eye position changes. To control for changes in the acoustic input to the eardrums brought about by the moving pinnae, we immobilized them for a final set of physiological recordings in one cat. We would have preferred to immobilize them reversibly by anesthetizing the branches of the facial nerve while holding a single neuron, but this proved too difficult. We also tried taping the pinnae, but this was unsuccessful in holding the pinnae steady. Finally, we resorted to acutely denervating the pinnae. Under sterile conditions, we cut the auriculoposterior and temporal branches of the facial nerve that innervate the caudal and rostral pinnae musculature, respectively. This procedure was successful in immobilizing the pinnae for about 3 or 4 wk, until the nerves regenerated, and presumably also deafferented the pinnae so that the proprioceptive signals were also severed.
Data analysis
In the psychophysical experiments, the dependent variable of interest was final eye position after saccadic eye movements to the acoustic targets. A criterion based on the variability of the velocity of the eyes during steady fixation was used to mark the start and end of an eye movement (Populin and Yin 1998b). Briefly, the end of and return to fixation were determined by the time at which the amplitude of the velocity function exceeded and returned to within 2 SD of the mean baseline velocity, respectively. The mean velocity baseline was computed from 100 ms before to 30 ms after target onset, a period during which the eyes were expected to be stationary. The eye position at the time of the return to fixation was considered to be the final eye position for a saccade. If corrective eye movements took place before the reward or within 500 ms after the end of fixation, the return to fixation measurement was carried out after the completion of the last movement.
Dot rasters and histograms were constructed to qualitatively evaluate neuronal responses. For quantitative evaluation, the mean number of action potentials per trial occurring within a predefined window was determined. For most neurons, a 500-ms window, starting 5 ms after the onset of the acoustic stimulus, was used. The auditory RF of a SC neuron is operationally defined as the area of space within which a stimulus evokes a response.
Because our primary concern was the role of eye position on the responses of SC neurons to acoustic stimuli, we usually began by studying the responses of the neurons to a probe from a single fixed position while varying the eye position using the sensory probe task. To determine if auditory responses were modulated by fixation position, 95% CIs were computed for the mean response from each fixation position. The auditory responses of a neuron were classified as modulated by fixation position if one pair of responses to different fixation positions was significantly different (P < 0.05). The exact shape of the resulting functions relating neural responses to fixation position is determined by the horizontal position of the speaker selected for studying the neuron, the shape of the RFs, and the way the RFs are modulated by fixation position.
When stable recording conditions and a cooperating animal coincided, a more complete representation of the neuron's RF was obtained by recording the responses evoked from sounds presented from different speakers along the horizontal axis for three to five different fixation positions. The data were plotted both as a function of the location of the speakers relative to the animal's head and as a function of the distance between the fixation point and the speakers (i.e., the motor error). The extent to which the RF functions obtained for the different fixation conditions shifted and became aligned or misaligned because of the change in coordinates constituted the test of the motor error hypothesis.
We computed the shift in azimuth of the RFs that resulted from two different eye positions relative to the eyes-centered condition as follows. For example, for each data point from the RFs measured when the cat fixated to the left, we measured how much that point was shifted in azimuth relative to the RF when the eyes were fixating straight ahead. This was done by computing at each data point the difference in azimuth at corresponding discharge rates, as shown by the dashed arrow in Fig. 2. This process was repeated using the data points from the RF obtained when the cat fixated at the center, as indicated by the solid horizontal arrow in Fig. 2. When the azimuth at which the line intersected the RF fell between data points, the azimuth of intersection was determined using linear interpolation. Shifts to the right of the center RF were positive, while shifts to the left were negative. These differences in azimuth were averaged. When measuring the differences, care was taken to respect the shapes of the RFs; for example, when RFs exhibited a "peak" response, the difference in azimuth took into account the medial and the lateral portions of the RFs to obtain a true estimate of the shift in the RFs. Neurons with RFs that were complex with many peaks and dips were not included. A motor error index (MEI) was computed by taking the ratio of the computed shift in the RFs and the size and direction of the shift in eye position. When the RFs exhibited a "peak" type profile, characterized by responses on either side of the peak that fell by 20% or more, an MEI was computed also for the medial edge of the RF only; this analysis is comparable to that performed by Jay and Sparks (1987). An MEI of 1.0 resulted if the RF shifted in the same direction and with the same magnitude as the shift in eye position. This process was repeated for RFs recorded with fixation to the right. For each neuron, we plotted the MEI for leftward fixations against the MEI for rightward fixations. Data points that fell in the upper right quadrant were therefore consistent with the motor error hypothesis, with perfect motor error for points at (1, 1).
After completion of the experiments, the cats were anesthetized with an intramuscular injection of ketamine and acepromazine, and two to three pins were placed in the brain with the recording microdrive at known positions within the recording chamber, rostrally and caudally to the recording area, and in parallel to the electrode penetrations. Figure 3 shows Cat6's midbrain with one pin inserted caudally to the left SC, and another pin inserted rostrally to the right SC. The hash patterns overlaying each SC represent the approximate areas in which electrode penetrations were made in relation to the location of the pins.
After placing the pins, a lethal dose of sodium pentobarbital (150 mg) was given intravenously, followed by perfusion through the heart with physiological saline and (10%) formalin. The brains were removed from the skull and stored in 30% sucrose at 4°C for 24–48 h. The midbrain was blocked using the pins as reference, frozen, cut into 50 μm sections in the coronal plane, and stained with cresyl violet. Individual penetrations were difficult to identify due to the long tenure of our cats, but numerous tracts traversing the SC could be observed. Camera lucida drawings and positional information referenced to the pins were used to confirm that the electrode penetrations reached the SC.
Effects of fixation position on sound localization accuracy
The first consideration was to determine whether Pöppel's (1973) paradox was behaviorally relevant, i.e., whether there was any effect of initial eye position on sound localization accuracy. To address this question we used the auditory standard-saccade task (Populin and Yin 1998b) with different initial fixation points (LED in Fig. 1A) and broadband noise stimuli as acoustic targets. Trials of this type were randomly intermixed among different types of visual and acoustic trials to numerous other targets (Populin and Yin 1998b). Figure 4 shows eye movements to an acoustic target at (0, 24°) recorded under those conditions. The cat was required to fixate LEDs at (±4, 0°) for 1,000 ms or an LED at (0, 0°) for 500 ms. The dots that form each trace represent digital samples of the analog output of the eye coil system. In ∼90% of acoustic trials on the horizontal plane, cats oriented with a single saccadic eye movement; for acoustic targets on the vertical plane, ∼95% of trials were completed with a single saccade. The slope of the main sequence computed from these eye movements was consistent with those measured in our previous work (Populin and Yin 1999). Data from six cats with different levels of experience were analyzed to address this question.
The trajectories of eye movements recorded in successful trials initiated from different fixation positions are plotted in Fig. 5, A–F. For the experiment shown in Fig. 5A, the acoustic target was located at (18, 13.5°), and the starting fixation positions were at (±9, 0°) and (0, 0°). Cat C9 was a novice subject, and the data were collected in the eighth behavioral session; the animal had not been exposed to this acoustic target until this session. As is typical of cats in head-restrained conditions (Populin and Yin 1998b), subject C9 undershot the acoustic target. Although the trajectories of the eye movements were different in the three fixation conditions, the eye movements ended up in roughly the same spatial location. The eyes took the most direct path when the fixation LED was on the same side of the target (9, 0°), and the most curved when the fixation LED was on the opposite side (−9, 0°).
The acoustic targets for the experiment shown in Fig. 5B were at (0, −22°) and (0, 24°) with fixation positions at (±3.5, 0°) and (0, 0°). Cat G1 was an experienced subject; these data, a subset of which is shown in Fig. 4, were collected in the 292nd behavioral session. This subject was very accurate in localizing sound sources regardless of initial eye position, and her data reveal no effect of initial eye position on sound localization.
The data shown in Fig. 5C are from subject C5's 11th experimental session. The acoustic target was located at (8, 13.5°). This was the first session in which subject C5 was presented with this acoustic target. The trajectories of the eye movements were similar to those of subject C9 in Fig. 5A. Although they show more corrections than the other subjects, they consistently reached the target.
For the experiments in Fig. 5, D and E, the acoustic target was located at (−18, 0°), and the initial starting positions were at (±9, 0°) and (0, 0°). Subjects C6 and C14 were well practiced at the time of the recordings, which were the 81st–83rd and 159th–163rd behavioral sessions, respectively. Both subjects mislocalized this acoustic target, although with different types of errors: while subject C6 localized the target below the horizontal plane, subject C14 localized it above. Interestingly, the pattern of errors made by each subject was consistent across the three experimental conditions.
Last, the data shown in Fig. 5F are from subject G7 and were collected in the 55th–60th behavioral sessions. The acoustic target was placed at (0, 24°), and the starting points were at (±6, 0°) and (0, 0°). Unlike the previous subjects, G7 exhibited a clear bias in its final eye position, which is indicative of incomplete compensation for the initial position of the eyes along the horizontal plane.
Figure 6 summarizes localization performance from the subjects presented in Fig. 5; data from some additional targets are included. In each panel, the position of acoustic targets is plotted with an asterisk, the starting fixation positions are plotted with filled symbols, and the corresponding mean final eye positions are plotted with open symbols of the same shape. Error bars represent 95% CIs computed for the x- and y-axes separately. All trials, successful and unsuccessful, were included in the analysis.
There was no effect of initial eye position on the accuracy with which subjects C9, G1, and C5 (Fig. 6, A–C) localized the acoustic targets. The 95% CIs computed for each of the mean final eye positions overlapped, indicating that they were statistically indistinguishable. Similar observations were made for targets at other positions.
Subjects C6's and C14's localization of an acoustic target at (−18, 0°) showed an effect of eye position, as revealed by the nonoverlapping 95% CIs computed for each of the three mean final eye positions. Subject C6's data (Fig. 6D) show a significant effect of eye position for the target at (−18, 0°) only. The magnitude and sign of the mean error for the eye movements initiated from the primary position (0, 0°) compared with those starting from fixation position to the right (9, 0°), but not to the left, of the midline are nearly proportional to their initial separation. There was no effect of initial eye position on the localization of other acoustic targets such as those located above (0, 9°) and to the right (18, 0°) of the primary position.
Subject C14 (Fig. 6E) made large, yet consistent, localization errors on both the horizontal and vertical components in all three conditions. However, the differences among the three mean final eye positions cannot be predicted based on the initial fixation position. Specifically, the final eye position for the (±9, 0°) starting fixation points was not significantly different. Furthermore, the localization error was smaller in the (9, 0°) than in the (0, 0°) fixation condition, which is the opposite of what one would expect if initial eye position was not taken into account to perform the eye movements to the acoustic target. Subject G7 (Fig. 6F) also showed a mixed pattern, i.e., an effect of initial eye position for some targets, those located along the vertical axis, but not an effect of initial eye position for others, those located along the horizontal axis.
Thus for the six cats tested, only one of them showed a consistent mislocalization that was related to initial eye position. The accuracy of saccades to auditory targets of three of the cats was unrelated to initial eye position, and for two cats, there was a small but inconsistent effect.
On average, the mean horizontal localization error computed from 267 trials from four cats, across three different initial eye positions, for targets with large horizontal components, was an undershoot of 35%.
Effects of fixation position on single SC neuron responses
As a microelectrode approached the SC from above, a predictable pattern of active and quiet areas was found in most penetrations. In cortex, we encountered visually driven activity followed by lack of activity in the superior cistern, which indicated that the SC was near. The signature of the superficial layers of the SC was intense neural activity that changed in rate as the cat moved its eyes or visualstimuli were swept across the screen in front of the cat. The onset of this pattern was abrupt and in marked contrast with the lack of activity just above. The area with this type of activity extended dorso-ventrally about 1.5 mm. The position of the tip of the electrode within the visual map of the SC (Feldon et al. 1970) was inferred from the responses evoked by stimulating discrete areas of the contralateral visual field with a visual stimulus while the cat fixated an LED at the primary position (0, 0°). We did not attempt to isolate neurons in the superficial layers of the SC. We searched for neurons that responded to acoustic stimuli as we lowered the electrode into the intermediate layers of the SC using a mixture of trials with visual and acoustic stimuli. In the caudal SC, auditory neurons were typically found between 1.5 and 3.5 mm ventral to the top of the superficial layers, but in approaching the rostral SC, the layer containing auditory neurons became progressively thinner (<0.5 mm), with fewer auditory neurons.
On isolating a neuron responsive to sound, we determined the stimulus position (from our available target locations) from which the strongest response could be evoked as well as the approximate threshold. The most sensitive speaker position was selected, and the level of the stimulus was set at ∼20 dB above threshold. Our first priority was to determine if the responses of the neuron using the sensory probe task, evoked from the most effective source location, were modulated by eye position. In cases in which we were able to maintain the recording for a prolonged period, we characterized other parts of the neuron's RF along the horizontal axis. As reported previously (Populin and Yin 2002), the responses of most SC auditory neurons to identical acoustic stimuli were smaller in the sensory probe task compared with the auditory fixation task. In some cases, the effect was so strong that there was virtually no response to the sensory probe.
With few exceptions, the 143 SC auditory neurons in our sample responded to broadband noise stimuli presented from sources in the contralateral field using the sensory probe task. Figure 7 shows the responses of a typical SC neuron to an acoustic sensory probe with the fixation point at (0, 0°). The cartoon at the center of Fig. 7 shows the position of the speakers relative to the fixation point. From top to bottom, Fig. 7A shows vertical and horizontal eye position plotted as a function of time, a dot raster showing the responses of the neuron in individual trials, and a histogram of the number of spikes/bin normalized by the number of trials, summarizing the responses of the neuron in the trials shown. The data are plotted synchronized to the onset of the acoustic stimulus at 0 ms. The eye position traces corresponding to the other targets were similar to those plotted in Fig. 7A, thus they were omitted for simplicity. This neuron, recorded from the right SC, responded most strongly to stimuli presented from the left hemifield (Fig. 7B). The response of this neuron to a long-duration (1,000 ms) broadband noise stimulus consisted of a burst of two to five action potentials with a mean first spike latency of 18.3 ms.
Modulation of SC auditory responses by eye position
If the auditory responses of SC neurons are modulated by eye position, it would indicate that auditory RFs are dynamic and that they might not encode the position of sound sources in head-centered coordinates. To address this issue, we used the sensory probe task (Fig. 1B) with different fixation positions and a single source of acoustic stimulation. Figure 8 shows responses of a neuron that was modulated by eye position. The three columns show responses of a neuron in the right SC to the same speaker at (−18, 0°), the area of space where the strongest responses could be evoked, while the cat was fixating three different positions: (±9, 0°) and (0, 0°). The largest responses (Fig. 8C) were recorded when the cat fixated an LED at (9, 0°), the farthest away from the sound source, and the smallest responses (Fig. 8A) were recorded when the cat fixated the LED closest (−9, 0°) to the sound source. Trials from each of the three fixation conditions were presented in random order, thus long-term changes in variability should be averaged out. The results are summarized in Fig. 8D, where mean spike count, normalized by the number of trials, is plotted as a function of horizontal fixation position. The error bars, representing 95% CIs, indicate that the mean responses recorded at the three fixation positions were significantly different. We studied 106 neurons in the SC using the test shown in Fig. 8 with a single speaker and three to five fixation positions, and three-fourths of them (81/106) were significantly modulated by eye position. Most of the neurons that were modulated by eye position were similar to that shown in Fig. 8, although we also found neurons whose largest responses occurred when the cat fixated the LED nearest the target.
Do RFs shift?
Figure 8 shows the responses of a neuron to acoustic stimuli presented from a single source recorded at three different eye positions. Ideally, one would like to extend these observations to a larger portion of a neuron's RF to determine if such changes in neural response were due to a shift in the RF as a function of eye position. Recordings of this type were possible in 28 neurons, 3 of which are shown in Fig. 9.
The motor error hypothesis predicts that auditory RFs will shift by an amount equal to the difference between the present eye position and the target, the motor error. Thus if a neuron encodes the location of sound sources in motor error coordinates, plotting the RFs as a function of motor error should make the RF profiles superimpose for different points of fixation. However, our RF profiles are undoubtedly incomplete because we did not sample in the caudal half of the auditory field, beyond 90° eccentricity, or in the vertical dimension. For example, the responses shown in Fig. 9A (left) are likely incomplete, because we only probed one flank of the RF. Nevertheless, the RFs of this neuron are better aligned when plotted as a function of motor error (Fig. 9A, right).
On the other hand, the RFs shown in Fig. 9, B and C, are more complete because we captured the peak and both flanks. Neuron C6-59 (Fig. 9B) had a narrow horizontal RF, which unlike most neurons, extended into ipsilateral space. The three functions, corresponding to the RFs measured in each of the three fixation conditions, are clearly separated at both the medial and lateral aspects of the field. Plotting the same data as a function of horizontal motor error aligns the three curves at both the medial and lateral edges, yielding a consistent measure of RF regardless of fixation position. RFs with similar profiles, although broader, were obtained from other neurons (e.g., Fig. 9C). The RFs of this neuron's medial edge were well aligned when plotted as a function of horizontal speaker position. However, unlike the neuron in Fig. 9B, plotting the same data as a function of horizontal motor error increased the separation between the curves.
To determine the extent to which RFs shifted as a function of fixation position, we computed a MEI as described in methods. The results of this analysis, taking into account all RF data available from each of the 28 neurons, are shown in Fig. 10A. For each neuron, the MEI due to leftward fixation is plotted against the MEI due to rightward fixation. Thus RF shifts consistent with the motor error hypothesis will fall in the upper right quadrant, with perfect motor error expected at an MEI of (1, 1). No shift would be indicated by an MEI of (0, 0). The mean MEI from all 28 neurons (0.65, 0.66), although smaller than (1, 1), is significantly different from (0, 0) because the 95% CIs do not encompass (0, 0). Thus the horizontal RFs of the neurons in our sample shifted in a direction consistent with the motor error hypothesis.
To compare our results with those of Jay and Sparks (1987), who measured the magnitude of auditory RF shift in the monkey by taking into account only the medial edge, we computed the MEI for only the medial edge of the RFs of our neurons as described in methods. The filled symbols in Fig. 10B show the MEI for the medial edge of neurons that had a peak type of RF (n = 16), as defined in methods, and the open symbols show the MEI for the remaining neurons (n = 13). The mean MEI from this analysis (0.81, 0.48) is also consistent with the motor error hypothesis, in agreement with the 54% compensation reported by Jay and Sparks (1987).
Potential contributions of the mobile pinnae
The modulation of auditory responses observed in the behaving cat appears to be similar to the modulation observed in the macaque monkey by Jay and Sparks (1987). Casual observation of the movements of the pinnae of both species indicates that cat's pinna movements are more prominent and thus more likely to change the acoustic input to the eardrum (Young et al. 1996), which can result in modulation of neuronal responses (Middlebrooks and Knudsen 1987).
Previously, we have documented that orienting behavior of cats with restrained heads to acoustic and visual targets involves stereotyped and coordinated movements of both the eyes and the pinnae (Populin and Yin 1998c). Because we required the cats to fixate on LEDs at different spatial locations to control eye position at the time of presentation of the acoustic stimuli, this would have systematically changed the position of the pinnae, which could have in turn altered the acoustic signals reaching the cat's eardrums. Such changes in acoustic input could account for the modulation of auditory responses in the SC documented above.
Figures 11 and 12 show the responses of two neurons, one which showed modulation by eye position and one which did not. The neuron in Fig. 11 displayed atypical responses because it was best driven by acoustic stimuli presented from an ipsilateral source (−18, 0°). We selected this neuron for illustration because the position of the left pinna was also recorded in the same trials. Five fixation positions were used: (±18, 0°), (±9, 0°), and (0, 0°). From top to bottom in Fig. 11A, each row of panels shows horizontal eye position, left horizontal pinna position, the responses of the neuron in individual trials in raster format, and a histogram normalized by the number of trials summarizing the neuron responses. As expected from our previous observations (Populin and Yin 1998c), the position of the left pinna at the time of stimulus presentation was a function of fixation position; a vertical arrow in each panel points to this portion of the pinna position records. The responses of neuron C6-75 to broadband noise stimuli presented from the same source at (−18, 0°) were significantly modulated by fixation position, and possibly, pinna position (Fig. 11B).
Figure 12 shows that the movements of the pinnae are not necessarily related to modulation of acoustic responses by eye position. The auditory responses of this unit were not modulated by fixation position, although the position of the left pinna at the time of presentation of the acoustic stimuli also varied as a function of fixation position. This neuron was recorded in the right SC, and the acoustic stimulus was presented from a target located at (−63, 0°).
The ear movement traces in Figs. 11 and 12 confirm that there were systematic changes in ear position related to the fixation of the LED at different spatial locations. Therefore there were also undoubtedly systematic changes in pinnae position during the collection of the data depicted in Figs. 8 and 9, each of which showed modulation with eye position. Thus the question arises as to whether the modulation with eye position was due to the change in acoustic input resulting from the movements of the pinnae or to an eye position signal that modulated the responsiveness of the neurons. On the other hand, the lack of modulation in Fig. 12 shows that changes in pinna position do not necessarily change the responsiveness.
Effects of pinna immobilization
The data presented in the previous section suggest the possibility that pinna movement associated with changes in fixation could account for the modulation observed in auditory responses of SC neurons. Changes in pinna position in the anesthetized cat have been shown to affect the spatial tuning of single SC neurons (Middlebrooks and Knudsen 1987). To see if eye position, independent of ear position, is a source of modulation of auditory responses in the SC of the behaving cat, we immobilized the pinnae of one cat by cutting the auriculoposterior and temporozygomatic branches of the facial nerve bilaterally and recorded the responses of single SC neurons to broadband acoustic stimuli.
Typical eye and pinna movement data recorded simultaneously with the auditory sensory probe task from the intact cat are shown in Figs. 11 and 12 before the immobilization surgery. At the time of these recordings, just before the pinnae immobilization procedure, subject C6's right pinna coil was broken, so baseline data only from the left pinna were acquired at the time. We chose not to submit subject C6 to an additional surgical procedure to repair the broken coil because baseline data from this structure, showing its movement, had already been recorded (Fig. 7 in Populin and Yin 1998c). The pinna adopted different positions during the period preceding the onset of the acoustic probe, corresponding to the different positions of the fixation LEDs. The largest pinna movement was recorded when the cat fixated the LED ipsilateral to the pinna, at (−18, 0°), and the smallest change was recorded when the cat fixated on the contralateral LEDs at (18, 0°) (Figs. 11 and 12A). Also consistent with our previous findings (Populin and Yin 1998c) are the short latency movement of subject C6's left pinna after the onset of the acoustic target.
Figure 13 shows that cutting the branches of the facial nerve was effective. After the pinnae immobilization procedure, C6's eyes continued to move normally (Fig. 13D), but there were no movements of either pinna when the cat fixated to the left or to the right (Fig. 13, C and E). Both the left and right pinnae remained motionless for ∼3 wk, after which time they began to recover their ability to move, presumably due to nerve regeneration. The single neuron recordings presented below were carried out while subject C6's pinnae were immobilized.
Physiological recordings after pinna immobilization
With the pinnae and head immobilized and the same acoustic stimuli presented while the cat fixated LEDs at different positions, any modulation of auditory responses in single SC neurons could not be attributed to changes in the acoustic input to the eardrum. Figure 14 shows the responses of a single neuron recorded from the left SC under those conditions. Broadband noise stimuli were presented from a speaker located at (63, 0°) at about 10–15 dB above threshold. The responses were transient, with little or no activity following the initial burst. Most importantly, they were significantly modulated by eye position (Fig. 14B). Most of the modulation was achieved within the initial burst. This neuron, like many in the deep and intermediate layers of the SC (Meredith and Stein 1983; Populin and Yin 2002), was bimodal and also responded to the visual presentation of the fixation LEDs when they were on the midline (Fig. 14A, middle) and in the contralateral hemifield (Fig. 14A, right). Arrowheads point to this transient activity in the histograms (Fig. 14A), which is synchronized to the presentation of the fixation LEDs, but because of the variable reaction time to make the saccade to the LED, they are not synchronized to the onset of the acoustic stimulus that was used to make the plots in Fig. 14.
We recorded from 24 neurons in a cat with immobilized pinnae, 20 of which were tested with only a single speaker. Approximately 3 wk after the immobilization procedure, some mobility returned to the pinnae; at this point, the experiment was terminated, and the brain tissue was recovered for histological examination. The auditory responses of all 20 neurons were modulated in the cat with immobilized pinnae.
We were able to characterize the horizontal component of the RFs of four neurons more extensively. Two examples of those recordings are shown in Fig. 15. In both cases, the RFs became more misaligned when plotted as a function of motor error. Figure 16A shows the MEIs based on all available RF data from these neurons, and Fig. 16B shows the MEIs computed only from the medial edge of the RFs. Data from one of the neurons were obtained at (±9, 0°) and (±18, 0°), which are plotted separately. Regardless of the type of analysis, none of the MEIs measured fell within the upper right quadrant, indicating that the neurons were modulated in a manner inconsistent with the motor error hypothesis. The mean and 95% CIs from the intact cat data (Fig. 10) are shown for comparison.
There are four major results from this study: 1) saccades made by cats to acoustic targets with their heads fixed are not generally affected by the initial position of the eyes; 2) most of the neurons in the deep and intermediate layers of the SC that are responsive to acoustic stimuli are modulated by eye position; 3) those neurons from which more complete data from the horizontal component of the RF are available show that the modulation is consistent with the predictions of the motor error hypothesis and account for approximately two-thirds of the necessary compensation; and 4) the modulation with eye position is not due to the change in pinna position with visual fixation since it is still present when the ears are immobilized. We will discuss each of these results in turn below.
Behavioral results
The behavioral data show that sound localization accuracy is generally not affected by the deviation of the eyes in the orbit at the time of acoustic stimulus presentation. This is not a surprising finding for grossly mislocalizing sounds simply because the eyes were not at the primary position would constitute a severe handicap for an animal. This means that the saccadic command to move the eyes must compensate for current eye position when a saccade is made to the same target from two different initial positions. For three of the six cats, there was no significant effect of initial eye position on the accuracy of saccades to targets, and for two other cats, there was a small but inconsistent effect. In only one cat (Figs. 5F and 6F) was there a consistent but small bias such that saccades starting from the left toward a vertical, but not a horizontal, target had a consistent error to the left, while those starting from the right had a rightward error. These results contradict Harris et al. (1980) in that they show that cats do compensate for deviations of the eyes from the primary position, but the results are in accord with both Hartline et al. (1995), who reported little effect of eye position on sound localization for deviations of about ±15°, and Peck et al. (1995), who reported partial compensation only. These data are also consistent with those from human (Yao and Peck 1997; Zahn et al. 1979) and monkey (Whittington et al. 1981) studies that found no effect of initial eye position on the accuracy of eye movements to acoustic targets.
Modulation of auditory responses in the deep and intermediate SC by eye position
The physiological data from the intact cat show that the responses of most SC auditory neurons (81/106) are modulated by eye position. In 28 neurons, we were able to sample the RFs more extensively along the horizontal axis using different points of fixation (Fig. 9). These data allowed us to examine the extent to which the horizontal component of the RFs of auditory neurons shifted as a function of fixation position. Based on the MEIs computed from the RFs, 68% of the neurons were modulated by fixation position in a direction consistent with the motor error hypothesis. The percentage increased to 76% when the MEI was based on the medial edge.
Since many neurons in the deep and intermediate layers of the SC are multimodal, responding to both visual and auditory stimuli (Meredith and Stein 1983), we should consider whether there is a visuomotor interaction that affects the eye position sensitivity shown in these experiments. Sixty-three percent of the neurons we studied were indeed bimodal (see example in Fig. 14 and Populin and Yin 2002). The example in Fig. 14 is typical of the short, transient visual response, which is over for almost 400 ms by the time the acoustic stimulus is turned on. While it is possible that there is some lingering inhibitory effect of the visual response on the auditory response many hundreds of milliseconds later, the parsimonious explanation would suggest that the effect is due to eye position and not to visio-motor interaction. Note that at the time the acoustic target is turned on, the eye is always fixating the LED, so there should be a constant visual input on every trial.
Thus our data show that 1) the responses of 76% (81/106) of auditory neurons are modulated by eye position, 2) the modulation of 68% (19/28) of neurons from which a more complete sampling of the RF is available is consistent with the motor error hypothesis, and 3) the mean MEIs, (0.65, 0.66) computed from all available RF data and (0.81, 0.48) for the MEIs computed from the medial edge of the horizontal RFs, account for a compensation of approximately two-thirds of the shift required for perfect motor error.
Because the cats undershot the targets significantly (Figs. 5 and 6), a more appropriate measure of the motor error would be the distance between the initial position of the eyes and the apparent location of the targets as indicated by the endpoint of the saccade. We refer to this as behavioral motor error (BME). Ideally, we would have preferred to compute the MEI for each neuron using BME and not the physical location of the targets relative to the head. Such MEIs would have accounted for a larger proportion of the deviation of the eyes from the primary position. Although we could estimate the BME from our psychophysical results (Figs. 5 and 6), we were unable to compute BME for the neurons we studied for the following two reasons. The oculomotor range (approximately ±20°) of the cat limits this analysis to neurons with RFs in a very small portion of the auditory field of most SC neurons, which extend out to 90° or more. Releasing the head to allow the cat to orient to targets anywhere in the frontal hemifield would have prevented us from systematically manipulating the position of the eyes in the head at the time of stimulus presentation of the acoustic stimuli. Despite these limitations, our results, computed as a function of target position, agree with those of Jay and Sparks (1987), who reported a compensation of 54% in the SC of the monkey and thus suggest that audio-visual interactions in monkey, a species with a larger oculomotor range, and cat SC are governed by similar rules. However, if we take into account the magnitude of the horizontal error made by our cats in localizing acoustic targets on the horizontal plane (Figs. 5 and 6), which on average was an underestimate of the targets by ∼35%, the mean MEIs computed from all available RF data (0.65, 0.66) actually account for the observed behavioral responses.
Our results suggest that the modulation with eye position seen in SC neurons correlates better with the actual motor response of the cat (undershoot of about 35%) than to the motor error. In the monkey, though there are data to suggest that the SC neurons are correlated better to motor error than to actual motor response (Frens and Van Opstal 1998; Stanford and Sparks 1994).
Effects of pinna immobilization on auditory responses in the deep and intermediate SC
Cats, however, move their pinnae in coordination with the eyes during the oculomotor tasks used in this study (Populin and Yin 1998c). Macaque monkeys make large pinna movements in response to unexpected sounds, but do not seem to hold the position for an extended time (Jay and Sparks 1987). The systematic changes in pinna position associated with fixating LEDs at different horizontal positions observed in our cats (Figs. 11 and 12) led us to consider that the modulation of responses to acoustic stimuli of SC neurons could be, at least in part, due to changes in the acoustic input to the eardrum.
To determine directly if the modulation of auditory responses in single SC neurons was due to the changes in the acoustics, we surgically immobilized the pinnae of one cat. Failure to observe modulation would have supported the hypothesis that acoustic factors were solely responsible for the modulation seen in the intact cat. Because all 24 neurons studied in the immobilized pinnae condition showed modulation by eye position, we reject the hypothesis that the modulation is caused exclusively by acoustic factors.
The patterns of the modulation of the spatial RFs documented in four neurons after immobilizing the pinnae, however, were inconsistent with our observations in the intact cat as assessed by the MEI (Figs. 10 and 16). It is possible that this discrepancy is due to the method by which we computed the MEI. As pointed out above, an MEI computed using the perceived target location rather than the actual location is likely a more appropriate measure of the motor error. Several factors precluded us from computing such a measure. First, as was the case in the experiments with intact cats, it was not possible to document this cat's localization accuracy at the range of eccentricities required to match those used to measure the auditory neurons' RFs. Second, due to the limited recording time under pinnae-immobilized conditions afforded by the regenerating facial nerves (∼3 wk), we were not able to conduct extensive psychophysical sound localization studies, focusing instead on the physiological recordings.
The data suggest that both an eye position signal and a pinna position signal contribute to shape motor error (Figs. 9 and 10). In the intact cat, copies of motor commands sent to the pinna may be used directly as a position signal or may be first evaluated against proprioceptive signals from the pinna. In either case, when the pinnae were immobilized, a mismatch would be expected between the desired position of the pinnae, the expected sensory consequences (i.e., acoustic filtering), and the eye position signal. While we are confident to have ruled out acoustical factors as the only source of the modulation, the small number of neurons recorded in the pinnae immobilized cat temper any conclusions drawn from the results following pinna paralysis.
Previous studies in the cat aimed at determining whether auditory RFs in SC neurons shift as a function of eye position produced conflicting reports. Harris et al. (1980) found no modulation in a sample of three neurons in the awake, untrained cat. They concluded that complex interactions between the visual and auditory representations in the SC are not needed because cats keep their eyes near the primary position most of the time, thus maintaining the alignment between the various sensory representations in the SC. Our results do not agree with this conclusion. Aside from their small sample of neurons, other methodological problems compromise their conclusions, including cats that were untrained, stimuli delivered without acoustic shielding, and eye positions that were averaged over large ranges.
The conclusions of more recent studies (Hartline et al. 1995; Peck et al. 1995) are consistent with our findings, but are difficult to evaluate. Hartline et al. (1995), for example 1) used essentially untrained cats, which made controlling eye position and distinguishing between sensory and motor responses problematic, 2) taped the pinna coils to the ears, which made them subject to slippage, 3) presented only a small amount of data, which made it difficult to evaluate their reliability, and 4) collapsed responses over relatively large ranges of eye and pinna positions. Peck et al. (1995) reported that their cats did not move their pinnae while performing their experimental tasks, although no records of pinna movements were presented. Our results (Figs. 11 and 12; Populin and Yin 1998c) show that cats with immobilized heads move their ears extensively and systematically in response to both visual and acoustic stimuli, even after thousands of trials. In agreement with our findings, Hartline et al. (1995) also reported that eyes and pinna tend to move in the same direction in a correlated fashion.
In summary, these results show the presence of an eye position signal that modulates the responses of SC neurons to acoustic stimuli. Eye positions signals in the SC are thought to originate from corollary discharges (Guthrie et al. 1983) and not from proprioception (Nelson et al. 1989). Several issues remain unexplained by our data and will require further experimental work. First, the nature of the modulation in the cat with immobilized pinnae is dissimilar to that in the intact cat. We suggest that pinna position, the motor command to move the pinnae or proprioceptive signals from the pinnae interact with eye position to modulate the SC cells, but further work is needed. Second, the responses of some SC neurons to acoustic stimuli were not modulated by eye position. While static RFs are inconsistent with the predictions of the motor error hypothesis, they can provide a head-centered reference against which signals from neurons with dynamic RFs can be compared. Third, the RFs of some neurons become more misaligned when plotted as a function of motor error. Such responses, which would appear to contradict the concept of motor error, might still fit the hypothesis if they were inhibitory to neurons that provide a gain modulation that varies with eye position. Fourth, as Jay and Sparks (1987) found in the SC of the monkey, the magnitude of the shift in RF observed in most neurons did not compensate for the deviation of the eyes from the primary position at the time of presentation of the acoustic stimuli. However, our data show that this undercompensation fits the undershooting saccades seen behaviorally. Behavioral motor error may be the important parameter in the SC.
This work was supported by National Institute on Deafness and Other Communication Disorders Grants DC-00116 and DC-02840 to T.C.T. Yin and DC-03693 to L. C. Populin and National Science Foundation Grant IBM-9904770 to L. C. Populin.
We thank J. Hudson and J. Sullivan for assistance, R. Kochhar, J. Sekulski, and S. Sholl for programming help, and R. Guillery for reading an earlier draft of this manuscript.
The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
- Copyright © 2004 by the American Physiological Society


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Skin Diseases of the Dog and Cat
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moreFollowing the super-success and critical acclaim for the First Edition, the new Second Edition of Skin Diseases of the Dog and Cat is being equally well accepted, by reviewers and readers alike. The strengths of the first edition have been preserved: - Problem-oriented approach; the book is arranged in twelve chapters covering dermatoses classified by their principle presenting sign. - Concise systematically structured text; definition, aetiology and pathogenesis, clinical features, differential diagnoses, diagnostic tests, management. Focus is on clinical practice and the need to explain the disease to the owner. - Superb and generously sized colour photographs and diagrams. - Clear user-friendly design - one condition per page or spread of pages. - Referenced thoughout to give the clinician access to the literature. Within the framework and in light of the exponential growth of information in small animal dermatology since first publication, the authors have revised the entire text for the new second edition, added twenty-one new sections and one hundred and thirty-one new colour photos. The result is a handbook that is practical, extensive and up-to-date in its content, beautifully illustrated and designed. The new second edition is of value to veterinary practitioners and students, veterinary nurses and technicans. less
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Title: Skin Diseases of the Dog and Cat
Author: Tim Nuttall; Richard G Harvey; Patrick J McKeever
Title: Skin Diseases of the Dog and Cat
Author: Tim Nuttall; Richard G Harvey; Patrick J McKeever
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- U.S. job growth surges in July, By Ana Swanson, August 4, 2017, Washington Post: "The U.S. economy added 209,000 jobs in July, according to government data released Friday morning, surpassing economists' expectations and suggesting the economy continues to thrive after an extended streak of job gains in recent years…"
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- 9 years after recession began, some states still unrecovered, By Jeff Amy (AP), May 18, 2017, St. Louis Post-Dispatch: "Call them the unrecovered — a handful of states where job markets, nine years later, are still struggling back to where they were before the recession. That's true in Mississippi, where job numbers and the overall size of the economy remain below 2008 levels. Unlike states that have long since sprinted ahead, Mississippi is struggling with slow economic growth and slipping population in a place that's rarely at peak economic health…"
Job Searching and the Unemployed – Illinois
More jobless Illinoisans are giving up the job search, study finds, By Alexia Elejalde-Ruiz, May 10, 2017, Chicago Tribune: "Though people often focus on unemployment rates as a measure of economic health, another telling data point is how many people are so discouraged with the job search that they're dropping out of the labor force altogether. A newly released survey found good news: Fewer unemployed Americans are giving up looking for work. But that's not the case in Illinois, where more people seem to be throwing up their hands…"
April 2017 US Unemployment Rate
- Economy adds strong 211,000 jobs, unemployment at 10-year low, By Paul Davidson, May 5, 2017, USA Today: "The labor market bounced back in April amid milder weather as employers added 211,000 jobs, providing evidence that weakness the prior month was a blip that likely won't keep the Federal Reserve from raising interest rates in June. The unemployment rate, which is calculated from a different survey, fell from 4.5% to 4.4%, a new 10-year low, the Labor Department said Friday…"
- Unemployment rate drops to lowest level in a decade in April as economy adds 211,000 jobs, By Ana Swanson, May 5, 2017, Washington Post: "The U.S. job market rebounded strongly last month and the unemployment rate fell to the lowest level seen in a decade, government data released Friday morning showed, calming fears that had bubbled up in the past month about the state of the economy…"
March 2017 US Unemployment Rate
- U.S. hiring slumped in March as employers added only 98,000 jobs, By Ana Swanson, April 7, 2017, Washington Post: "The momentum in the U.S. labor market flagged in March, new government data showed Friday, with the private sector and the government adding only 98,000 jobs, the lowest gain in nearly a year, as winter storms weighed on economic activity…"
- Job growth loses steam as U.S. adds 98,000 in March, By Nelson D. Schwartz, April 7, 2017, New York Times: "Job growth turned in a disappointing showing in March, according to data released on Friday by the Labor Department. It is the latest official snapshot of the state of the American economy…"
February 2017 US Unemployment Rate
- U.S. added 235,000 jobs in February; unemployment rate dropped to 4.7 percent, By Ana Swanson, March 10, 2017, Washington Post: "The U.S. economy added a healthy 235,000 jobs in February, according to government data released Friday morning, surpassing economists' expectations and likely clearing the way for the Federal Reserve to raise interest rates this month. The unemployment rate ticked down to 4.7 percent, compared with 4.8 percent in January, and wages rose by 6 cents to $26.09 in February, after a 5-cent increase the month before…"
- U.S. gains a healthy 235,000 jobs in February; unemployment rate falls to 4.7%, By Don Lee, March 10, 2017, Los Angeles Times: "Warm weather and rising business optimism helped the U.S. economy to create another burst of job growth last month, giving President Trump an early confidence boost and all but assuring that the Federal Reserve will nudge up interest rates next week. Employers added 235,000 jobs in February, about as many as in January and well above analysts' expectations and the average monthly payroll growth for all of last year, the Labor Department said Friday…"
Unemployment Benefit Programs – Missouri, Ohio
- Missouri lawmakers vote to slash unemployment benefits, By Austin Huguelet, February 23, 2017, St. Louis Post-Dispatch: "The Missouri House on Wednesday reaffirmed its commitment to a plan that would cut the amount of time people can claim jobless benefits to one of the shortest periods in the country…"
- Unemployment fraud still costing state millions, By Cornelius Frolik, February 21, 2017, Dayton Daily News: "Ohio has significantly reduced fraudulent overpayments to people collecting unemployment insurance since instituting measures designed to identify and recover money from benefit cheats…"
The US Structurally Unemployed
The new face of American unemployment, By Jeanna Smialek and Patricia Laya, February 7, 2017, Bloomberg: "Even at so-called full employment, some 20 million Americans are left behind. They're looking for work, out of the labor force but unhappy about it, or report working part-time when they'd prefer more hours, according to data released last week. Their plight comes even as the U.S. flirts with what economists consider the maximum level of employment for the first time since before the recession, having added 15.8 million jobs since the start of 2010. While some of America's jobless are simply between gigs, those persistently stuck out of work are called the structurally unemployed…"
January 2017 US Unemployment Rate
U.S. starts year with job surge, but pay gains are weak, By Patricia Cohen, February 3, 2017, New York Times: "Job growth was quick out of the gate in the new year, the government reported on Friday, as employers added a healthy 227,000 workers to their payrolls in January. But despite a surge of local minimum-wage increases in states across the country, wage growth was meager. The official jobless rate rose slightly, to 4.8 percent, but for good cause: More people were lured back into the work force…"
December 2016 US Unemployment Rate
U.S. economy creates a modest 156,000 jobs in December, By Don Lee, January 6, 2017, Los Angeles Times: "Job growth slowed at the end of last year while the unemployment rate ticked higher — signs that the tight labor market is resulting in diminished hiring even as it is pushing wages higher. The U.S. economy added 156,000 net new jobs last month, down from an upwardly revised 204,000 created in November, the Labor Department said Friday…"
Basic Income – Finland
Finland trials basic income for unemployed, By Jon Henley, January 3, 2017, The Guardian: "Finland has become the first country in Europe to pay its unemployed citizens an unconditional monthly sum, in a social experiment that will be watched around the world amid gathering interest in the idea of a universal basic income. Under the two-year, nationwide pilot scheme, which began on 1 January, 2,000 unemployed Finns aged 25 to 58 will receive a guaranteed sum of €560 (£475). The income will replace their existing social benefits and will be paid even if they find work…"
Unemployment System – Ohio
No fix this year for troubled unemployment system, By Jessie Balmert, December 7, 2016, Cincinnati Enquirer: "Ohio lawmakers won't overhaul the state's troubled unemployment system – at least not until April, they say. After days of furious negotiating, lawmakers, business and union leaders came up short of a comprehensive fix that would please both the people who receive unemployment benefits and the employers who pay for them. Instead, lawmakers said Tuesday, they plan to freeze unemployment benefits for 2018 and 2019. Employers currently pay taxes on their employees' annual wages up to $9,000, and lawmakers plan to increase that to $9,500. The national average is $13,407…"
November 2016 US Unemployment Rate
U.S. economy added 178,000 jobs in November; unemployment rate dropped to 4.6 percent, By Ana Swanson, December 2, 2016, Washington Post: "The U.S. economy added 178,000 jobs in November, while the unemployment rate fell to 4.6 percent, a level not seen since August 2007, according to government data released Friday morning. The first employment report since voters went to the polls last month shows an economy in strong shape as President-elect Donald Trump prepares to take office…"
October 2016 US Unemployment Rate
Final jobs report before election shows promising wage growth, By Don Lee, November 4, 2016, Los Angeles Times: "The last monthly jobs report before Tuesday's presidential election offered some encouraging news for workers: Job growth remains steady and pay is rising at a faster rate. Employers in October added 161,000 jobs, a little less than analysts' average forecast but still a solid pace consistent with a healthy labor market. That's more than enough to absorb new entrants to the workforce and keep the jobless rate from rising…"
US Unemployment Claims
U.S. unemployment claims at lowest level since 1973, October 13, 2016, Denver Post: "The number of Americans applying for first-time unemployment benefits held at a four-decade low last week, a sign employers are holding on to workers as the labor market tightens. Initial jobless claims, a proxy for layoffs, were unchanged at 246,000 in the week ended Oct. 8, the Labor Department said Thursday. The latest two weeks registered at the lowest level since November 1973…"
Unemployment Benefits – North Carolina
$2 billion in trust for jobless benefits. Is it time to increase unemployment checks?, By Richard Craver, October 6, 2016, Winston-Salem Journal: "North Carolina has more than $2 billion in its trust fund to pay unemployment benefits, a level close to what federal guidelines suggest for reserves. State Division of Employment Security officials told legislators Wednesday the amount should be enough for the agency to handle the payout demands of the next recession without having to borrow again from the federal government. However, some unemployment advocacy groups say the trust fund should be doubled to at least $4.2 billion before state officials should be secure with the amount…"
Youth Unemployment – Chicago, IL
Chicago tackles youth unemployment as it wrestles with its consequences, By Alexia Elejalde-Ruiz, September 1, 2016, Chicago Tribune: "Margo Strotter, who runs a busy sandwich shop in Chicago's Bronzeville neighborhood, makes it a point to hire people with 'blemishes.' But young people? She sighs and shakes her head. They often lack 'the fundamental stuff' — arriving on time, ironing their shirts, communicating well, taking direction — she said. She doesn't have time to train workers in the basics, and worries she's not alone. 'We are going to wind up with a whole group of people in their 40s and 50s who can't function,' said Strotter, owner of Ain't She Sweet Cafe. As Chicago tackles what some have termed a crisis of youth joblessness, it must reckon with the consequences of a failure to invest in its low-income neighborhoods and the people who live there. There aren't enough jobs, and the young people vying for them are frequently woefully unprepared because of gaps in their schooling and upbringing. The system has pushed them to the back of the hiring line…"
June 2016 State Unemployment Rates
18 states see significant job gains, but unemployment rises, By Josh Boak (AP), July 22, 2016, Albany Times Union: "Unemployment rates ticked up noticeably in six states in June, even as employers continued to add jobs and the hiring outlook improved. The unemployment rate in 21 states is now significantly below the national figure of 4.9 percent, while it's higher in 14 states…"
Unemployment Insurance – North Carolina
North Carolina ranks among top-five worst states for the jobless, By Richard Craver, June 20, 2016, Winston Salem-Journal: "North Carolina's inclusion among the five worst states for unemployment insurance beneficiaries may be a source of shame or pride, depending on the value placed upon the drastic cuts that went into effect in July 2013. The state was ranked 46th by research firm 24/7 Wall St. in a report released June 11. Louisiana is listed as the worst state, followed by Alabama, Mississippi and Alaska. In a report released Thursday, North Carolina ranked 49th in terms of what percentage of UI applicants receive benefits at 12.4 percent for 2015. That study comes from the Center for American Progress, Georgetown Center on Poverty and Inequality, and National Employment Law Project…"
Internet Access and Unemployment – Detroit, MI
Unemployed Detroit residents are trapped by a digital divide, By Cecilia Kang, May 22, 2016, New York Times: "In downtown Detroit, start-ups and luxury retailers are opening up and new office buildings are being built as the city works to recover from its deep economic problems. Six miles to the north, in the neighborhood of Hope Village, residents like Eric Hill are trying to participate in that progress but are running into hurdles. His difficulties were apparent on a recent Tuesday when he entered a crowded public library to use the computers to look for a new job. With no Internet service at home or on his mobile phone, Mr. Hill had few options to search work listings or file online job applications after losing his stocking job at a pharmacy five months ago…"


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Behind the Success of Bayyinah: A lesson for Muslim Entrepreneurs
I started teaching Arabic courses and khutbahs here and there and more and more my heart wasn't in my career. The short courses I was teaching once in a while at local masjids were surprisingly successful. I was more surprised by them than anyone else. When I did decide to take the leap of faith and go full time into teaching Arabic as a career which would fund my own studies of the Qur'an, those closest to me were all skeptical. They had good reason to be. But I was confident in my plan, my 'business' plan had been attacked left and right by my close friends in order to further refine my strategy and by the grace of Allah, I dove in.
The success of Bayyinah and my own popularity online is what most people see. What my closest friends and family know all too well are the toil, labour, sacrifice and difficulty that were all a big part of making this thing work. I look back and I wonder how I and my family survived the ordeal of the first few years.
I'll just say that I have tremendous respect for socially responsible entrepreneurship. That's what I consider myself. I know that a lot of times our youth see a success story and say to themselves. 'If he can do it, why can't I?' Though I respect the enthusiasm and appreciate the sentiment, I want my young brothers and sisters to understand that I had never worked harder in my life until I started working for myself. I used to think working 40-50 hours a week with full time college was tough.
When I started this project, those days seemed like a vacation! I'm grateful that I delivered newspapers in the dead of the winter in Queens NY when I was in high school, that I ran around in a shoe store helping customers try out different sizes, that I got yelled at everyday for messing up one thing or another at a travel agency…the list goes on and on. I'm happy that I was responsible for my own finances from an early age because it gave me respect for work and a sense of responsibility.
I didn't go into full time Islamic work with delusions of grandeur and fame. I just wanted to teach and provide a valuable service. I honestly didn't expect the popularity that Alhamdulillah I and our organization have enjoyed. He is the one who gives honor and He is the one who can take it away at any moment. But I do want to underscore that the help of Allah comes to those who remain committed, serious, optimistic and most importantly sincere.
May Allah count us all among them.
Interview conducted by HOTD


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Upgrade of WP 2.0 RC-1
Just wanted to tell you about a successful upgrade from WP 1.5.2 to WP 2.0 RC-1.
System: My blog resides at an ISP ( web hotel ) running Apache and MySQL. This is how it went:
1. Unpacked the distribution locally.
2. Copied the database connection strings from wp-config.php to wp-config-example.php and saved it as the new wp-config.php.
3. Deleting everything from my blog except the themes and plugins I've added.
4. Uploaded the new WordPress bunch to the server
5. Reached for the /wp-admin/upgrade.php from my browser
6. Clicked the link Upgrade WordPress>>
( After getting an introductory error message "WordPress database error: [Table 'petitpub.wp_usermeta' doesn't exist] – SELECT meta_key, meta_value FROM wp_usermeta WHERE user_id = '1'" )
That's it!
It all worked like a breeze, including my themes and user and my permalinks.
I'm happy 🙂
- The topic 'Upgrade of WP 2.0 RC-1' is closed to new replies.


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Slick's Que Co. based in Sisters Oregon combines authentic pit BBQ, signature rubs and sauces, a festive environment and great service to create the most flavorful and mouthwatering experience you'll ever have in a fun, family-friendly and fast casual restaurant.
Walk in the door of Slick's Que Co. and you'll immediately feel welcome as part of our extended family; you'll see, hear, smell and taste the heart and soul that we put into our food and guest experience. Owner and pitmaster Roy Slicker has traveled all over the country visiting BBQ joints large and small, has served on the Board of Directors, and is a past President of the National BBQ Association, all with the goal of serving the most tender and flavorful "Que" you'll ever taste.
In June of 2010 and again in 2015 and 2016, Slick's Que Co.was honored as one of "The Best of the Best BBQ Restaurants in America" by the BBQ News Publication. Slick's Que Co. is quickly becoming a popular destinations for people all over the West, with most of our guests making the trek as a result of the continuous word-of-mouth referrals about the food, service and atmosphere.
Read testimonials from our visitors.
© 2017 Slick's Que Co. All rights reserved


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Reference Number. 170099
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Uxbridge Glaziers
Uxbridge Glaziers was founded to provide clients with a complete glass and glazing solution at competitive costs. From initial discussions and design meetings through to site surveys and final handover, we aim to provide clients with complete confidence and reassurance based on years of glazing experience. (Our senior management team has a combined experience of over 30 years serving the glazing industry! ) With a strong commitment to customer service and attention to detail, Uxbridge Glaziers prides itself on being one of the few glazing contractors capable of meeting the diverse requirements of today?s glazing markets.149 Grosvenor Crescent
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Grants and Funding: Extramural Programs (EP)
NLM Extramural Programs (EP)
EP Overview
The Extramural Programs (EP) Division of the National Library of Medicine (NLM) offers grants for research projects and research training in biomedical informatics and data science. Biomedical informatics research applies computer and information sciences to improve the use of biomedical information. Read More
EP News Archive
- Trans-NIH Strategic Plan for Research on Women's Health (NOT-OD-17-108)
- Reminder: NIH Applications Must Be Complete and Compliant With NIH Policy and Application Instructions At Time Of Submission (NOT-OD-17-105)
- Notice of Assistance Avaliable to Institutions Impacted by Hurricane Harvey (NOT-OD-17-111)
- Notice of Change to NIH Policy for Issuing Certificates of Confidentiality (NOT-OD-17-109)
- Policy Supporting the Next Generation Researchers Initiative (NOT-OD-17-101)
- NIH Request for Information (RFI): Pediatric research that All of Us may be uniquely positioned to enable (NOT-PM-17-004)
- Reminder: NIH Natural Disaster Policy - Hurricane Harvey (NOT-OD-17-106)
NLM Research Spotlight Archive PECASE Awards
NLM Lecture Series Archive
Research Funding
Career Development
- Biomedical Informatics Research Training (T15)
- Information for Training Directors
- NIH Pathway to Independence (K99/R00)
- NLM Career Development Award in Biomedical Informatics and Data Science (K01)
- Ruth L. Kirschstein NRSA Individual Predoctoral Fellowships (F30/F31)
- Research Grant Supplement (Diversity)
- Research Grant Supplement (Reentry)


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the evening of October 7th at 6:30 PM the CCMS Band was honored at the
new Clay City Hall. A proclamation was presented to honor the band for
past and present accomplishments.
The Clay-Chalkville Middle School Band Receives Recognition
The Clay-Chalkville
Middle School Band has been invited to perform as a feature ensemble at
the Southeastern United States Concert Band Clinic. The invitation came
this summer from the Seus Board of Directors asking the band to perfom
on Friday, December 6, 2013. The band will perform for an audience of
over 400 middle school band students and directors from Floriday,
Georgia, Mississippi, Tennessee, Arkansas, and Alabama. Alsin
attendance at the concert will be several university and pre-service
music educators, as well as internationally known composer, Quincy
Only two middle school
bands throughout the Southeastern United States are invited each year.
The band is under the direction of Dr. Chris J. Walker and Ms. Erin