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Recent studies have suggested that the basal ganglia are related to motivational control of behavior. To study how motivational signals modulate motor signals in the basal ganglia, we examined activity of midbrain dopamine (DA) neurons and caudate (CD) projection neurons while monkeys were performing a one-direction-rewarded version (1DR) of memory-guided saccade task. The cue stimulus indicated the goal position for an upcoming saccade and the presence or absence of reward after the trial. Among four monkeys we studied, three were sensitive to reward such that saccade velocity was significantly higher in the rewarded trials than in the nonrewarded trials; one monkey was insensitive to reward. In the reward-sensitive monkeys, both DA and CD neurons responded differentially to reward-indicating and no-reward-indicating cues. Thus DA neurons responded with excitation to a reward-indicating cue and with inhibition to a no-reward-indicating cue. A group of CD neurons responded to the cue in their response fields (mostly contralateral) and the cue response was usually enhanced when it indicated reward. In the reward-insensitive monkey, DA neurons showed no response to the cue, while the cue responses of CD neurons were not modulated by reward. Many CD neurons in the reward-sensitive monkeys, but not the reward-insensitive monkey, showed precue activity. These results suggest that DA neurons, with their connection to CD neurons, modulate the spatially selective signals in CD neurons in the reward-predicting manner and CD neurons in turn modulate saccade parameters with their polysynaptic connections to the oculomotor brain stem.
The basal ganglia play an important role in voluntary saccadic eye movements (Hikosaka et al. 2000). This is done mainly by serial inhibitory connections: one from the caudate (CD) to the substantia nigra pars reticulata (SNr) and the other from the SNr to the superior colliculus (SC). Spatial information necessary for saccades may be provided by the inputs from the frontal eye field and the surrounding prefrontal cortices (Selemon and Goldman-Rakic 1985; Yeterian and Pandya 1991; Parthasarathy et al. 1992). Recent studies from our laboratory indicated that this basal ganglia mechanism modifies a saccade depending on whether the saccade is followed by reward (Kawagoe et al. 1998; Sato and Hikosaka 2002; Takikawa et al. 2002a,b). An important question is: Where does the reward-related information originate? A candidate for the input would be midbrain dopaminergic neurons that are located in and around the substantia nigra pars compacta (SNc) (Andén et al. 1964). Indeed, animal experiments provided some hints that DA neurons are related to reward and motivation (Robbins and Everitt 1996). DA neurons in monkeys respond to reward and to sensory stimuli that predict reward (Schultz 1998). Specifically, DA neurons respond to unexpected reward and show depression of activity when reward is unexpectedly omitted (Schultz et al. 1997) or shifted (Hollerman and Schultz 1998). Other lines of evidence suggest a common role of DA neurons in animals and humans. Thus reinforcement of brain self-stimulation in animals and drug addiction in humans seem to be mediated by DA neurons (Wise 1996). Human imaging studies have suggested that rewarding experiences are associated with changes in neural activity (Delgado et al. 2000; Knutson et al. 2000) or increases in DA transmission in the striatum (Koepp et al. 1998). These observations provided experimental evidence for recent theories of reinforcement learning (Barto 1994; Houk et al. 1995; Montague et al. 1996).
Despite a large body of data on DA neurons, it is still unclear how their signals are used to modify motor outputs. A likely site where DA neurons may influence motor signals is the dorsal striatum (including the CD and putamen), since a majority of DA neurons project to the striatum (Lynd-Balta and Haber 1994) and its dorsal part is thought to have motor functions (Alexander and Crutcher 1990). Inside the striatum are cellular and molecular mechanisms with which DA neurons can exert strong influences on striatal projection neurons (Calabresi et al. 1997; Nicola et al. 2000). DA depletion in the unilateral CD induces deficits in initiation of saccades to the contralateral side (Kato et al. 1995; Kori et al. 1995) or contralateral hemineglect (Miyashita et al. 1995). Since CD projection neurons are a major origin of eye movement signals in the basal ganglia (Hikosaka et al. 2000), DA neurons are in a good position to induce the reward-contingent modulation of activity of CD neurons and other basal ganglia neurons.
Using a modified memory-guided saccade task (called 1DR task) (Kawagoe et al. 1998), in which reward was given for one particular direction out of four targets, we found that activity of CD neurons is strongly modulated by the presence and absence of an upcoming reward (Kawagoe et al. 1998) and the position at which reward is given (Lauwereyns et al. 2002b; Takikawa et al. 2002a). To study the function of midbrain DA neurons in reward-oriented saccades, we recorded from CD neurons and DA neurons in the same monkeys using 1DR. We found that activity of DA neurons was even more clearly modulated by an upcoming reward. Interestingly, one monkey showed no reward-dependent modulation of saccades, unlike the other monkeys. In this monkey neither CD neurons nor DA neurons exhibited reward-dependent modulation of visual cue responses. Such individual differences were consistent with the hypothesis that the reward dependencies of DA neurons and CD neurons are causally related such that saccades are modulated depending on an upcoming reward (Takikawa et al. 2002b).
We used four male Japanese monkeys (Macaca fuscata): monkeys G, H, K, and M. Under general anesthesia, we implanted a head holder, chambers for unit recording, and a scleral search coil (Hikosaka et al. 1993). All surgical and experimental protocols were approved by the Juntendo University Animal Care and Use Committee and are in accordance with the National Institutes of Health Guide for the Care and Use of Animals. The monkeys were trained to perform saccade tasks, especially a memory-guided saccade task (Hikosaka and Wurtz 1983b). Eye movements were recorded using the search coil method. We recorded extracellular spike activity of presumed projection neurons in the CD in all four monkeys (G, H, K, and M) and presumed DA neurons in and around the SNc in three monkeys (G, H, and M).
Task procedures
The monkeys performed the memory-guided saccade task in two different reward conditions: all-directions-rewarded condition (ADR) and one-direction-rewarded condition (1DR). We used a set of four target locations of equal eccentricity (either 10° or 20°), arranged in either normal or oblique angles, depending on the neuron's response field. For every neuron recorded, we required the monkeys to perform one block of ADR and four blocks of 1DR (i.e., four different rewarded directions).
A task trial started with the onset of a central fixation point that the monkeys had to fixate (see Fig. 1). A cue (spot of light, duration: 100 ms) came on 1 s after onset of the fixation point at one of the four target locations, and the monkeys had to remember its location. After a random period (1-1.5 s), the fixation point turned off, and the monkeys were required to make a saccade to the previously cued location. The target came on 400 ms later for 150 ms at the cued location. The saccade was judged to be correct if the eye position was within a window around the target (usually within 3°) when the target turned off. The correct saccade was indicated by a tone stimulus. The next trial started after an intertrial interval of 3.5-4 s.
In ADR, every correct saccade was rewarded with a liquid reward together with the tone. In 1DR only one of the four locations was rewarded while the other directions were not rewarded (as illustrated in Fig. 1). The rewarded direction was fixed in each block of experiments which consisted of 60 successful trials. Even for the nonrewarded direction, the monkeys had to make a correct saccade. If the saccade was incorrect, the same direction was repeated. The target cue was chosen pseudorandomly such that the four directions were randomized in every subblock of four trials; thus one block of experiment (60 trials) contained 15 trials for each direction. 1DR was performed in four blocks, in each of which a different direction was rewarded (highly rewarded for monkey K). Other than the actual reward, no indication was given to the monkeys as to which direction was currently rewarded. The average amount of reward in one block of experiment was set approximately the same between 1DR and ADR; in other words, the amount of reward in a rewarded trial of 1DR was approximately four times larger than the amount of reward in a trial of ADR.
The monkeys were first trained to perform the ADR task. They were then trained to perform the 1DR task. Before the recording experiment started, the monkeys had been trained on the 1DR task extensively for more than 300 blocks.
Recording procedures
Eye movements were recorded using the search coil method (Enzanshi Kogyo MEL-20U) (Judge et al. 1980; Matsumura et al. 1992; Robinson 1963). Eye positions were sampled at 500 Hz. The behavioral tasks as well as storage and display of data were controlled by a computer (PC 9801RA, NEC, Tokyo, Japan). The unitary action potentials were passed through a window discriminator (Bak INC, Model DDIS-1), and the times of their occurrences were stored with a resolution of 1 ms.
Before the single-unit recording experiment, we obtained MR images (Hitachi, AIRIS, 0.3T) such that they were perpendicular to the recording chamber. We then determined the recording sites in the substantia nigra based on the chamber-based coordinates (Kawagoe et al. 1998). The recording sites were further verified by MR-imaging a plastic guide tube through which the electrodes were inserted.
Single-unit recordings were performed using tungsten electrodes (diameter: 0.25 mm, 1-5 MΩ, measured at 1 KHz, Frederick Haer). To introduce the electrode into the brain, we first inserted a stainless steel guide tube (OD 0.85 mm, ID 0.60 mm) containing the electrode. A hydraulic microdrive (Narishige, MO95-S) was used both to insert the guide tube and subsequently to advance the electrode into the brain. We sometimes implanted a plastic guide tube (OD 1.1 mm, ID 0.8 mm) semipermanently at the location where DA neurons were concentrated. The location of the guide tube was visualized on MRIs and was confirmed to be directed to the SNc.
Identification of DA neurons and CD projection neurons
We searched for DA neurons in and around the SNc (François et al. 1985). Guided by MRIs, we first searched for the SNr, where neurons showed high-frequency spontaneous firing (Hikosaka and Wurtz 1983a). The identity of the SNr was assured if such rapid firing stopped in relation to saccade tasks (Sato and Hikosaka 2002). Often intermingled with or just medial to SNr neurons (usually ≤2 mm) were presumed DA neurons that fired irregularly and tonically at about 5 spikes/s with broad spike potentials. Extracellular spikes may have an initial positive component or may be followed by prolonged positive component (Schultz and Romo 1987). A neuron with these features was thus determined to be a DA neuron. We then examined whether it responded to the delivery of reward. A drop of water as a reward was given with a random time interval (4-9 s) while the monkey was sitting in the dim experimental room. If the neuron responded to reward, we had the monkeys perform at least one block of ADR and four blocks of 1DR (i.e., four different rewarded directions). In addition, we sometimes repeated 1DR blocks to confirm the reproducibility of the neuron's behavior.
We recorded extracellular spike activity of presumed projection neurons, which showed very low spontaneous activity (<1 Hz) (Hikosaka et al. 1989a), not of presumed interneurons, which showed irregular tonic discharge (Aosaki et al. 1994). To find CD projection neurons, we let the monkeys perform 1DR continuously because CD neurons were often very quiet if the monkey was not performing a task. If a CD projection neuron was found and it fired at any period while the monkey was performing 1DR, we had the monkeys to perform at least one block of ADR and four blocks of 1DR.
Data analysis
determination of a response. We analyzed the postcue response of CD neurons and DA neurons and the postreward response of DA neurons. For each response, we determined its presence by comparing the spike frequency in the control period and the spike frequency in the test period using the Wilcoxon signed-rank test (P < 0.001) separately for rewarded and nonrewarded trials. For CD neurons two kinds of test period after the cue stimulus were set for each neuron: 200-500 ms for visual response and 500-1000 ms for sustained response. If the neuron showed the postcue response significantly at 200-500 and/or 500-1000 ms, we determined that a postcue response was present. We focused on reward enhanced type whose test activity was higher in rewarded trial than nonrewarded trial. For DA neurons the test period for the postcue response was 100-300 ms (for excitatory responses) and 200-500 ms (for inhibitory responses). The control period was set just before the onset of the fixation point for both of CD and DA neurons, its duration being the same as that of the test period. The test period for the response to reward was set to be 100-400 ms after the time when reward would be delivered. We then determined the presence of the reward response in the same way as the cue response (Wilcoxon signed-rank test, P < 0.001).
statistical analysis for the reward and spatial selectivity. A normalized cue response was calculated, for each trial, as the spike frequency during the test period minus the spike frequency during the control period. We performed nonparametric analysis for reward and spatial selectivity for the four-block data of 1DR for the neuron's normalized response. The first eight trials in each block were excluded from the analysis. Reward selectivity of the postcue response in 1DR was tested by the Mann-Whitney U test (P < 0.01) by comparing the responses in the rewarded trials and the responses in the nonrewarded trials. Spatial selectivity of the postcue response was determined by Kruskal-Wallis test (P < 0.01) by comparing the responses in four cued directions in 1DR. We used nonparametric tests because in many trials there was no spike in a test or control period and hence the sampled spike frequencies did not show the normal distribution.
reward/nonreward discrimination of saccade parameters (see fig. 2 and table 1). For each experiment including the four blocks of 1DR, we obtained mean values of saccade parameters (velocity, latency, and amplitude) separately for rewarded trials (52 trials: 13 trials × 4 blocks) and nonrewarded trials (156 trials: 39 trials × 4 blocks). The first eight trials in each block were excluded from the analysis. We then performed a statistical comparison for the entire set of experiments between the rewarded and nonrewarded conditions. For this purpose, we used the Wilcoxon signed-rank test (P < 0.0001) by pairing the mean value of rewarded trials and the mean value of nonrewarded trials for each experiment.
reward/nonreward discrimination of error rate (table 1). For each experiment including the four blocks of 1DR, we obtained the number of errors separately for rewarded and nonrewarded trials. We then calculated total error rate across experiments.
After recording was completed, monkeys H, G, and M were anesthetized with an overdose of pentobarbital sodium and perfused through the heart with 4% paraformaldehyde. The brain was blocked and equilibrated with 20% sucrose. Frozen sections were cut every 50 μm in the planes parallel to the electrode penetrations so that complete tracks were visible in single sections. The sections were stained with cresyl violet. Reconstruction of the recording sites was based on microlesions (5 μA for 180 s) made at the end of some recording experiments. Other recording sites were estimated based on these microlesions. To examine the dopaminergic activity in the brain of monkeys G and M, we used tyrosine hydroxylase (TH) immunohistochemistry to visualize dopaminergic neurons in and around the SNc and axon terminals in the CD, according to the procedure described previously (Kato et al. 1995). One of every five sections was subject to TH immunohistochemistry and this was done for the anterior-posterior dimension including the entire substantia nigra.
Behavioral sensitivity to reward
We trained four monkeys to perform 1DR of the memory-guided saccade task. Before the recording experiment started, all monkeys had been trained for the 1DR task for more than 300 blocks. As shown in Fig. 2A, in three monkeys (H, G, and K), the peak saccadic velocity was significantly faster when the saccade was followed by reward (ordinate) than when the saccade was not followed by reward (abscissa) (Wilcoxon signed-rank test, P < 0.0001, also see Table 1). This phenomenon has been reported previously (Takikawa et al. 2002b). In monkey M (Fig. 2B), however, the peak saccadic velocity was not different between the two conditions (Wilcoxon signed-rank test, P > 0.01, Table 1). In monkeys H and K, the saccade latency was significantly shorter in rewarded than in nonrewarded trials (Wilcoxon signed-rank test, P < 0.0001, Table 1). Furthermore, the coefficients of variation of saccade parameters were significantly lower in monkeys H, G, and K in rewarded trials than in nonrewarded trials (Wilcoxon signed-rank test, P < 0.0001, parenthesis in Table 1). The error rate was significantly lower in monkeys H, G, and K in rewarded trials than in nonrewarded trials (Wilcoxon signed-rank test, P < 0.0001, Table 1). These tendencies were absent in monkey M. On the other hand, the saccade amplitude was not significantly changed (Table 1). Therefore we judged that three monkeys (H, G, and K) were sensitive and one monkey (M) was insensitive to an upcoming reward. The saccade velocity was generally higher in monkey M than the other monkeys regardless of reward condition (Table 1).
Search for DA neurons and CD projection neurons
A total of 368 neurons in three monkeys (H, G, and M) were determined to be DA neurons (see methods). Among them, 203 (55%) responded to the delivery of reward by phasically increasing its discharge rate. We further examined these reward-responsive DA neurons using the ADR and 1DR. The other reward-unresponsive neurons were not examined further, although we cannot exclude the possibility that they also were DA neurons. The data presented in this paper are based on 71 DA neurons for which we were able to examine at least one block of ADR and four blocks of 1DR. They were recorded from monkeys H (n = 16), G (n = 32), and M (n = 23).
A total of 397 neurons in four monkeys (H, G, K, and M) were determined to be CD projection neurons (see methods) and were related to the performance of the memory-guided saccade task. Among them, 293 CD neurons responded to the visual cue stimulus in the memory-guided saccade task (Kawagoe et al. 1998); other types of neurons (Lauwereyns et al. 2002a; Takikawa et al. 2002a) are not included if they showed no visual response. The data presented in this paper are based on 206 CD neurons for which we were able to examine at least one block of ADR and four blocks of 1DR. They were recorded from monkeys H (n = 84), G (n = 59), K (n = 36), and M (n = 27).
The behavioral sensitivity (indicated by the saccade velocity) to reward was well correlated with the neuronal sensitivity to reward. In the three reward-sensitive monkeys (H, G, and K), a majority of both CD and DA neurons tended to be sensitive to an upcoming reward. In the reward-insensitive monkey (M) a majority of CD and DA neurons were insensitive to reward. In the following we mainly concentrate on three monkeys in which we recorded from both DA neurons and CD neurons (monkeys H, G, and M).
Comparison of neuronal activity between reward-sensitive and reward-insensitive monkeys
Figure 3 shows typical examples of a DA neuron (top) and a CD neuron (bottom) recorded in reward-sensitive monkey H. The data represent the neuronal sensitivity to an upcoming reward. The DA neuron responded to reward with a burst of spikes when the reward was delivered while the monkey was not engaged in a task (data not shown). However, when the same reward was delivered in response to the correct performance of the monkey in ADR or 1DR, DA neurons showed no discernable response except for the first few rewarded trials (Fig. 3, top, shaded parts). Instead, they responded to the cue stimulus in a selective manner in 1DR. In the first block of 1DR in which the right-up (RU) direction was rewarded (RU column), the DA neuron exhibited a burst of spikes after RU cue that indicated an upcoming reward. In the second block in which the left down (LD) direction was rewarded (LD column), the neuron responded to LD cue with a similar burst of spikes, but no longer responded to RU cue. In this way, the DA neuron changed its response completely such that it always responded to the cue that indicated the upcoming reward (Wilcoxon signed-rank test, P < 0.0001). The magnitude of the cue response was invariable whichever cue was rewarded (Kruskal-Wallis test, P = 0.6105). For the other three directions that indicated no reward, the neuron showed some suppression in activity (Wilcoxon signed-rank test, P < 0.0001). These results indicate that DA neurons acquired a particular position-reward association very quickly and flexibly. In contrast, the neuron showed no sign of response in ADR in which reward was delivered in every trial (ALL column).
The CD neuron in this monkey (Fig. 3, bottom) also showed a remarkable flexibility. In ADR the neuron responded strongly to the contralateral (RU and RD) cue stimuli. The neuron changed its preferred direction completely in 1DR. It responded most strongly to whichever cue stimulus indicated an upcoming reward. This kind of reward contingency was a common feature among CD projection neurons (Kawagoe et al. 1998). On the other hand, the CD neuron showed no reward response (Fig. 3, bottom, shaded parts).
We found that neuronal activity was remarkably different in the reward-insensitive monkey M (Fig. 4). The DA neuron shown in Fig. 4, top, did not respond to the cue stimulus in either 1DR or ADR, but responded with a burst to reward itself. This pattern was completely opposite to the DA neuron in the reward-sensitive monkeys (Fig. 3, top).
The CD neuron (Fig. 4, bottom) responded selectively and vigorously to RU cue stimulus in ADR and 1DR (RU row). Unlike the CD neuron in the reward-sensitive monkey H, the postcue response appeared unchanged depending on the rewarded direction. An analysis using Mann-Whitney U test showed no significant difference between the rewarded trials (RU column) and the unrewarded trials (LU, LD, and RD columns) (P = 0.9505) for RU cue direction in 1DR. Also unlike in the reward-sensitive monkey H, the neuron's preferred direction RU was ipsilateral to the recorded neuron.
The difference in neuronal activity between the reward-sensitive and reward-insensitive monkeys is summarized as population histograms in Fig. 5. DA neurons in the two reward-sensitive monkeys (H and G) were very similar (Fig. 5A, left). While the response to reward was negligible (not shown), the response to the cue stimulus was robust and selective for the expected outcome of reward. The response to the reward-indicating cue (thick black line) appeared as a single positive peak that reached about 16 spikes/s in both monkeys. The response to the nonreward-indicating cue (thick gray line) consisted of an initial small positive peak that was followed by a delayed negative peak, again in both monkeys. On the other hand, DA neurons in reward-insensitive monkey M were completely different as shown in Fig. 5B, left. The cue response was very weak and did not discriminate between the reward-indicating and nonreward-indicating cues.
Among 48 DA neurons in the reward-sensitive monkeys, 42 neurons showed significant cue responses (Wilcoxon signed-rank test, P < 0.001, Table 2). All cue-responsive DA neurons showed reward-dependent modulation (Mann-Whitney U test, P < 0.01, Table 2). In contrast, among 23 DA neurons in the reward-insensitive monkey M, only 1 showed significant cue response, and even that response was not modulated by an upcoming reward (Table 2). Instead, most DA neurons (17/23) in monkey M continued to respond to reward delivery during 1DR, while the reward response in reward-sensitive monkeys were less (Table 2). Thus the DA neuronal responses during 1DR in the reward-sensitive monkeys were quite different from those in the reward-insensitive monkey. On the other hand, basic characteristics of DA neurons were similar. The spontaneous discharge rate was 4.5 ± 1.2 (monkey H), 4.9 ± 1.2 (monkey G), and 4.7 ± 1.1 (monkey M) (mean ± SD) spikes/s. More than half of DA neurons responded to reward when it was delivered without warning: 21/33 (64%) (monkey H), 125/237 (53%) (monkey G), and 57/98 (58%) (monkey M).
CD neurons also showed a clear contrast. Population activities of CD neurons of reward-enhanced type are shown in Fig. 5. Anticipatory precue activity was evident in the reward-sensitive monkeys (H and G) (Fig. 5A, right), not in the reward-insensitive monkey M (Fig. 5B, right). The postcue response was equally frequent (Table 3) and was spatially selective in both types of monkeys (Kruskal-Wallis test, P < 0.01, Table 3). As reported previously (Kawagoe et al. 1998) reward selectivity occurred in the reward-sensitive monkeys (Fig. 5A, right), but not in the reward-insensitive monkey M (Fig. 5B, right). A majority of postcue-responsive CD neurons in the reward-sensitive monkeys (104/127, 82%) showed significant reward-dependent modulation of the postcue response (Mann-Whitney U test, P < 0.01, Table 3). Among them 76 neurons showed stronger responses to the reward-indicating cue than to the no-reward-indicating cue (positive reward modulation) as shown in Fig. 5A, whereas in 28 neurons the response was stronger to the no-reward-indicating cue (negative reward modulation). In the reward-insensitive monkey M, only 2 of 20 neurons showed reward-dependent modulation (both positive) (Table 3).
Comparison of DA and CD neuronal activity in reward-sensitive monkeys
The data shown so far indicate that the reward-dependent activity in DA neurons and CD neurons exists in the reward-sensitive monkeys, but not in the reward-insensitive monkey. This raises the possibility that DA and CD neuronal responses are either causally related or share common causes. To study the relationship between DA and CD neurons in the reward-sensitive monkeys, we examined spatial and temporal factors of the neuronal responses (Fig. 6). In Fig. 6A the DA and CD cue responses in the reward-sensitive monkey H are shown separately for the contralateral cue (left) and the ipsilateral cue (right). Only 4 of 42 DA neurons showed significant contralateral versus ipsilateral spatial selectivity (Kruskal-Wallis test, P < 0.01 Table 2). In contrast, CD neurons responded better to the contralateral cue than to the ipsilateral cue provided that it indicated an upcoming reward (Fig. 6). A majority of CD neurons (82/127 in the reward-sensitive monkeys and 16/20 in the reward-insensitive monkey) showed significant spatial selectivity (Table 3). These data indicate that CD neurons receive spatially selective inputs that do not originate from DA neurons.
Figure 6B shows changes in DA and CD neuronal activity within a block of 1DR task. In the first subblock of four trials (value 1 in the abscissa) in which the cue was presented in all four directions, both DA and CD neurons showed no differential responses to the reward-indicating cue (thick black) and the no-reward-indicating cues (thick gray). This was natural because the monkey knew the rewarded direction only by experiencing the trials in different directions. DA neurons and CD neurons were different in this initial cue response: while DA neurons showed little response to the contralateral or ipsilateral cue (little departure from the background activity), CD neurons already showed full responses that were stronger to the contralateral cues than the ipsilateral cues. This again indicates that the spatially selective responses of CD neurons originate from neurons other than DA neurons.
In the second subblock or thereafter, both DA and CD neurons differentiated their cue responses depending on the presence and absence of the upcoming reward. In DA neurons, the response to the reward-indicating cue became positive (i.e., increase from the background activity) while the response to the no-reward-indicating cue became negative (i.e., decrease from the background activity). In CD neurons, the response to the reward-indicating cue increased slightly while the response to the no-reward-indicating cue decreased monotonically.
The data in Fig. 6B suggested that the activity of DA neurons and the activity of CD neurons covaried within one block of 1DR. To illustrate their relationship we replotted the time-course data in Fig. 6B for monkey H as a scatterplot in Fig. 7, but now for monkey G and M as well. Each data point indicates the mean cue response of DA neurons (abscissa) and the mean cue response of CD neurons (ordinate). Only the responses to the contralateral cue are shown. In the reward-sensitive monkeys (H and G) there was a clear positive correlation between the mean cue responses of DA and CD neurons (Spearman ρ = 0.869 P < 0.0001, ρ = 0.840 P < 0.0001, respectively). If we consider the CD cue response as a function of the DA cue response, any positive response of DA neurons (compared with the background activity) would result in a positive shift of CD responses (compared with their initial responses in the first subblock); any negative response of DA neurons would result in a negative shift of CD responses. No such relationships were found in the reward-insensitive monkey M (Spearman ρ = -0.317, P > 0.03), possibly because the dynamic range of DA cue responses was negligible.
General behavior
All four monkeys were healthy and had a good appetite throughout the experiments. However, monkey M was remarkably different from the other three monkeys. When humans showed up in front of their home cages, most monkeys would show some signs of affection, aggression, obedience, or withdrawal. Monkey M showed no such sign. When an investigator tried to touch them, most monkeys would avoid the touch. Only when an intimate relationship has been established, monkeys may invite tactile contacts by presenting themselves toward a human mate. Monkey M made no such sign of avoidance or invitation, remaining expressionless. When presented with a favorite food such as banana, most monkeys would grab it and eat it hastily. Monkey M showed no sign of an immediate interest; he would not reach for the food for a while, although he would eat it eventually. A monkey usually discriminated humans, showing emotional attachments with the investigator who worked with the monkey. Monkey M showed no sign of such social discrimination. Although monkey M was usually indifferent to human observers, he was more sensitive to a sudden sensory stimulus, especially sound, by orienting to the source of the stimulus.
Interestingly, monkey M was able to learn saccade tasks, including the memory-guided saccade task, as quickly and accurately as the other monkeys. He even had no difficulty in learning 1DR. Most monkeys were reluctant to, and occasionally refused to, make a saccade to the nonrewarded direction. Monkey M showed no sign of reluctance. However, monkey M tended to get drowsy during experiments, a tendency so robust as to be unusual.
Recording sites of DA neurons
After making more than 20 electrode penetrations to the region around the SNc for each monkey, we could determine the region where DA-like neurons were recorded, on which the subsequent recordings were centered. Near the end of the recording session, we selected representative locations for electrode penetration and, when DA neurons were recorded, we made electrolytic microlesions at the recording sites. Histological examination revealed that the microlesions were located in the region inside or just mediodorsal to the SNc. This was true for both the reward-sensitive monkeys, G (shown in Fig. 8, A-D) and H (not shown), and the reward-insensitive monkey M (Fig. 8, F and H-J). By aligning the sections that contained the microlesions on the adjacent TH-immunostained sections, we found that the microlesions and reconstructed recording sites were among TH-positive neurons that were most likely to be DA neurons for both monkey G (Fig. 8E) and monkey M (Fig. 8G). The lack of behavioral and neuronal sensitivity to the upcoming reward in monkey M raised the possibility that the DA mechanism of monkey M may be abnormal. However, TH immunohistochemistry showed no gross abnormalities.
Relationship between CD neuronal activity and saccade behavior
In the memory-guided saccade task, a saccade-motor instruction is given by a positional cue stimulus. Many CD neurons respond preferentially to the contralateral one (Hikosaka et al. 1989b). By using a one-direction version of the memory-guided saccade task (1DR), we previously found that the CD postcue response was strongly modulated by reward condition (Kawagoe et al. 1998). Typically, the postcue response was much larger when the cue indicated an upcoming reward. 1DR task also gave rise to a robust behavioral outcome: saccades to the rewarded direction were faster and earlier than saccades to the nonrewarded directions (Takikawa et al. 2002b). These observations suggested that CD neurons create the reward-dependent modulation of saccade parameters. This idea was supported by a recent study indicating that CD neuronal activity, including the postcue response, is positively correlated with the velocity of contralateral saccades (Itoh et al. 2003). How then might DA neurons contribute to the neuronal and behavioral outcomes of reward?
Relation between DA neurons and CD projection neurons
There were similarities and differences between DA neurons and CD projection neurons. They were similar in that their cue responses were stronger when the cue indicated an upcoming reward in 1DR than when the cue indicated no reward (positive reward modulation). Whereas this rule applies to all DA neurons, a small fraction of CD neurons showed the opposite reward dependency (negative reward modulation, data not shown). Distinct differences were found in their task specificity, spatial selectivity, and precue activity. DA neurons showed postcue response in 1DR exclusively, whereas CD neurons responded both in ADR and 1DR. DA neurons showed no spatial preference, whereas CD projection neurons usually responded to the contralateral cue rather than to the ipsilateral cue. The spatial preference of CD neurons was present also in ADR. Precue activity was evident in CD neurons (Takikawa et al. 2002a), but not in DA neurons, while postcue activity were evident in both CD and DA neurons. To summarize, the cue response of DA neurons seems to be determined exclusively by the reward contingencies (not by the spatial localizations), whereas that of CD projection neurons is determined by both the reward contingency and spatial localization.
If DA neurons project to the CD and make synaptic connections, the reward contingency for the cue response of CD neurons may, at least partly, be derived from DA neurons. Indeed, the within-block development of DA cue responses was similar to that of CD cue responses.
These results imply that the spatial factor of CD cue responses is derived from non-DA neurons. One plausible candidate would be the inputs from the cerebral cortex, although other inputs such as those from the thalamus (Matsumoto et al. 2001) could also contribute. Cortical inputs to the CD that carry spatial attention or working memory would include the frontal eye field, supplementary eye field, intraparietal sulcus area, and dorsolateral prefrontal cortex (Parthasarathy et al. 1992; Selemon and Goldman-Rakic 1985; Yeterian and Pandya 1991). These cortical signals carrying spatial information may be modulated by DA signals carrying reward information. The modulation would usually be positive because in a majority of CD neurons the cue response increases when reward is expected (Kawagoe et al. 1998). Such a facilitatory effect of DA neurons may be mediated by D1 receptors (Hernández-López et al. 1997; Reynolds et al. 2001). In the small number of CD neurons for which the cue response is depressed if reward is expected (Kawagoe et al. 1998), the inhibitory effect of DA neurons may be mediated by D2 receptors (Hernández-López et al. 2000; West and Grace 2002). In any case, it is plausible to think that the interaction of the spatial signals from the cerebral cortex and the reward-related signals from DA neurons in CD projection neurons are the reason why the monkeys behaved (or made saccades) in a reward-sensitive manner. In monkey M, DA neurons showed no response to the cue stimulus in 1DR, and under our hypothesis CD neurons could not acquire the ability to discriminate between the reward-indicating and no-reward-indicating cues.
Although we favor the hypothesis above, we cannot rule out the possibility that the relationship is reversed: from CD to DA. A main target of CD neurons is the SNr (François et al. 1994; Hedreen and DeLong 1991; Parent and Hazrati 1994). Many SNr neurons send recurrent collateral axons to DA neurons in and around the SNc (Tepper et al. 1995). Since CD projection neurons and SNr projection neurons are both GABAergic, spike activity of CD neurons would lead to disinhibition of DA neurons. Under this alternative, postcue activity of CD neurons might lead to or facilitate postcue activity of DA neurons.
Precue anticipatory activity
The precue anticipatory activity was present in many CD neurons in the reward-sensitive monkeys, but not in the reward-insensitive monkey M. It is unlikely that the precue activity affected saccade parameters directly, because the activity occurred before the motor instruction. Nonetheless, the precue activity might influence the saccade parameters indirectly. As illustrated in previous studies from our laboratory (Lauwereyns et al. 2002a; Takikawa et al. 2002a), the precue activity was present selectively when a particular direction, especially a direction contralateral to the recorded neuron, was rewarded. Many of the CD neurons with precue activity also showed postcue responses with similar direction selectivity. Note, however, the meaning of the direction selectivity was different: selectivity for rewarded direction for precue activity; selectivity for instructed direction for postcue response. This combination of precue activity and postcue activity gave rise to selective boosting of the postcue response when the preferred direction was rewarded. The enhancement appeared as a positive bias or boosting rather than a gain change of the postcue response (Lauwereyns et al. 2002a). The enhanced postcue response would then facilitate saccades.
Our recent studies have suggested that the precue activity could indeed contribute to changes in saccade parameters. When the monkey was instructed to make a saccade to the cue stimulus on its appearance, the precue activity was followed by a saccade immediately (Lauwereyns et al. 2002b). When a contralateral side was rewarded, the precue activity was higher and the saccade latency was shorter. The result is consistent with the hypothesis that stronger precue activity leads to facilitation of a saccade to the contralateral direction. Monkey M would certainly lack the hypothetical facilitatory effects of CD precue activity on contralateral saccades. This might partly account for the lack of reward-dependent modulation of saccades in monkey M.
Possible pathological mechanisms
Monkey M's behavior was clearly different from other monkeys that we have tested. He rarely showed motivated behavior, emotional reactions, or any sign of affect. And yet, monkey M was more sensitive to sudden sensory stimuli than other monkeys. We, the authors, agreed that these behaviors were pathological, likened to human psychiatric or neurological disorders. These behavioral symptoms were somewhat similar to parkinsonism, although monkey M showed no obvious sign of motor deficits. Parkinsonian patients often have poor expression (called "mask-like expression"), like monkey M; they are often reluctant to initiate an action spontaneously (hypokinesia), like monkey M; they often showed reflex blink hyperexcitability (see Basso et al. 1996), as monkey M continued to respond to sudden sensory stimuli. The similarities suggested that the basal ganglia of monkey M did not function normally. In this monkey, DA neurons showed almost no response to the cue stimulus while CD neurons were not influenced by expected reward, unlike the other monkeys. One obvious difference between parkinsonian patients and monkey M was that DA neurons become scarce in parkinsonian patients, but were present and active in monkey M. Histological examination suggested that there were plenty of TH-immunoreactive neurons in the substantia nigra, which probably correspond to DA neurons. Despite the apparently normal chemical feature, these DA neurons did not function normally. It is possible that CD neurons recorded in monkey M may have not acquired the ability to predict an upcoming reward, perhaps due to the lack of reward-predicting signal in DA inputs. Alternatively, we may have recorded from different types of CD neurons in these monkeys. However, the latter explanation seems unlikely because we rarely found reward-predicting responses in CD neurons in monkey M.
On the other hand, monkey M's behavior was somewhat similar to symptoms of schizophrenia or autism. Schizophrenic and autistic patients are often alienated and lack emotional or social contact with other people, like monkey M. In fact, it has long been suspected that DA dysfunctions are involved in schizophrenic symptoms (Weinberger 1987). However, pathophysiology of schizophrenia remains largely elusive. It is largely because there has practically been no primate model of schizophrenia. One way to solve this problem would be to rely on the natural incidence of behavioral disorders, but the probability of finding such behavioral disorders would be very low. One way to circumvent this difficulty would be to collect animals with behavioral disorders from research institutions world wide. Researchers would probably be willing to give away such "psychiatric" animals since they are usually inappropriate for most experiments. The collection would be a precious data base for research on disordered mind and behavior.
We thank K. Nakamura, H. Nakahara, J. Lauwereyns, and B. Richmond for helpful comments, H. Itoh for analyzing data, and M. Kato for designing the computer programs.
This work was supported by a Grant-in-Aid for Scientific Research on Priority Areas (C) of Ministry of Education, Culture, Sports, Science, and Technology; Core Research for Evolutional Science and Technology of Japan Science and Technology Corporation; and Japan Society for the Promotion of Science Research for the Future program.
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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Five members of the Shabak Israel Security Agency were wounded, Monday morning, when a military jeep in which they were travelling was rammed by a terrorist from a village near the Samarian (Shomron) Palestinian Authority city of Tulkarm, in the area of the Samarian Arab village of Deir Sharaf, between the Jewish communities of Einav and Shavei Shomron.
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Robin Brownlee
August 15 2011 01:39PM
For those of you who care what The Hockey News thinks, the scribes over there are picking the Edmonton Oilers to finish dead-last in the Western Conference in 2011-12.
While that doesn't necessarily mean the staff at THN is predicting a third consecutive 30th place finish for the Oilers -- we don't get a breakdown of overall NHL standings in their predictions in the item -- writers at the publication, at one time considered the Hockey Bible, obviously don't see Edmonton improving enough to escape the conference basement.
The item reads:
"Why: Ryan Nugent-Hopkins was added to the exciting core of youth in Edmonton, but like the Senators, the Oilers still have a couple years of bumps and bruises ahead of them. While the prospects should have fans giddy with excitement, the fact is Edmonton is buried in a deep conference full of Stanley Cup contenders, so expectations have to be tempered. Once their prospects earn some experience, the Oilers will be off and running."
There's also a link highlighting off-season changes forecaster.thehockeynews.com/hockeynews/hockey/teamhome.cgi
Truth be told, I have difficulty mustering much outrage regarding THN's prediction, probably because I don't think they're far off in terms of how the Western Conference stacks up.
At 25-42-12 for 62 points last season, the Oilers obviously have a long way to climb. Last season, Colorado was 14th with 68 points and Columbus finished 13th with 81 points.
So, even allowing for a 20-point improvement in the standings, and that's hardly a given, realistically, how many teams in the West are the Oilers likely to pass in the pecking order? One? Two?
While I think a 20-point improvement is possible -- if Ryan Whitney and Ales Hemsky play something approaching full seasons and if Ryan Smyth, Cam Barker, Ben Eager and Eric Belanger represent the upgrades I think they do -- it's not going to vault the Oilers many places up the standings.
My best guess is the Oilers will overtake more teams from the Eastern Conference in overall standings, bottom-feeders like Ottawa, the New York Islanders and Florida Panthers, than they do teams from the West.
Still weeks from training camp, my first inclination is they finish right on the cusp of a third straight lottery pick, or 24-26th overall.
Listen to Robin Brownlee Wednesdays and Thursdays from 3 p.m. to 5 p.m. on the Jason Gregor Show on TEAM 1260.
Oilers will finish 6th in the West in 2011-12; a 37 point improvement over last year! Islanders also crack the top eight in the east
August 16 2011, 12:44AM
We are forgetting that we had a terrible injury situation last year which helped immensly in our quest for the first draft pick, If we have a relatively injury-free year I should think our standings will be around 21-23 area. There is no way if our team had been healthy last year we would have finished last.
The Oilers were terrible before guys started getting injured.
They were on pace for 60 points (last place!)30 games in before they started racking up injuries.
Last in the West is not an unreasonable prediction given that this team, even before injuries, was so awful last year. We were outplayed in every (yes, EVERY) aspect of the game, especially the physical part. Jones was the only forward who made a consistent effort to battle along the boards and go to the net for screens and rebounds. Too many "skill" players tried to score via fancy stickhandling and were repeatedly knocked off the puck, only to slowly skate back to their own end while the opposition had a quality 3-on-2 the other way. Gagner and Cogs couldn't win a faceoff or backcheck to save their lives. Omark's diminuitive size makes his efforts fruitless. Gilbert is a spectator in his own end. The coaching staff believe Khabby is a starting goalie. This all has to change.
I think the addition of Smyth and Belanger is crucial to changing this lousy approach to the game, because the coaching staff and other veterans have forgotten how to win, and team management and the fans have forgotten what quality hockey looks like. Until management stops using draft pedigree as their main criteria for acquiring veterans and then drafting their buddies' kids as our future (Gagner, Musil, Simpson?), this team will perpetually rely on top 5 picks to get out of the basement. We lucked out by having Smyth demand to come back here, because Lowe and Tambellini are as big a gong show as Garth Snow and the NY Islanders.
Sorry for the rant, but I don't like losing and I'm tired of posters here being enamoured with how many first round "prospects" this team is collecting. Who cares where our players were drafted, just win some GD games!
Smyth is going to help this team a lot. His point totals over the last couple of seasons should not be discounted. The PP will be much better and it all starts with having a player willing to muck it up in front of the net. Smyth is a huge upgrade on Penner when you consider consistancy and ability.
The D will be more physical and a little faster.
To me this adds up to 9-11 in the West. I think this season will look a lot like either of the 06-07/07-08 season. Meandering a few spots out from the playoffs and winning at around 40% clip. The good news is that the team looks like its on the rise rather than just hanging on. The team will have some good rental players that can be traded at the deadline for picks which can then be packaged for an NHL goalie next summer.
@Mason Storm
With Florida minus Vokoun they are toast.
Florida's top 6F: Fleischmann, Booth, Upshall, Weiss, Versteeg, Kopecky. Top 4 D, Campbell, Jovocop, Kulikov, Weaver and Gubranson playing this year.
They are weaker in goal, but much stronger everywhere else.
Toronto are relying on Reimer far too much and will implode.
Agreed that TO has a chance of being a lottery team
The Islanders always screw everything up They have as much young talent as the Oilers and are older. If Dipietro hits the IR early they will not be bottom 5
Phoenix is finished without Bryzgalov. They were a playoff team last year and LaBarbara is a fine young goalie. They lost some talent, but are not a lottery team.
New Jersey will continue their decline, especially after they trade Parise.
You are talking about the team with the best record in the NHL after they fired their coach. Way off.
Ottawa are in the midst of a re-build and plan look like they'll bottom out
Agreed, but Anderson will have them winning more games than they should.
Dallas minus Richards can't compete in the Pacific.
They still have a solid top 6F and but poor top 4D. If Lethonen plays decent they are not a lottery team. If he doesn't they may be in the lottery
Calgary are a year older and a year worse
Calgary had the 13th best goal differential in the NHL last year, and best goal differential of any team that didn't make the playoffs. If Kipper plays ok they have a shot at the playoffs. They are not a lottery team.
I think you underestimate just how crappy the Oilers are in term of goal differential (best predictor of winning hockey games) and how little they did to improve.
August 16 2011, 08:15AM
Florida's D isnt that good. Gudbransen for all his hype, will be eaten alive against NHL competition, especially if he plays top 4 minutes. Jovocop is getting older and Campbell has absolutely nothing to play for.
And never bet against Garth Snow screwing it all up.
August 16 2011, 08:21AM
Florida's D isnt that good. Gudbransen for all his hype, will be eaten alive against NHL competition, especially if he plays top 4 minutes. Jovocop is getting older and Campbell has absolutely nothing to play for.
And never bet against Garth Snow screwing it all up.
I see it mentioned daily, maybe we could see a list of all of Snow's "screw ups".
Florida's D isnt that good. Gudbransen for all his hype, will be eaten alive against NHL competition, especially if he plays top 4 minutes. Jovocop is getting older and Campbell has absolutely nothing to play for.
And never bet against Garth Snow screwing it all up.
Florida does have a good D.
I forgot to include Garrison into the top 4 D.
Garrison and Weaver handled the toughs last year and did an amazing job considering the team they played for.
See this link: http://blogs.thescore.com/nhl/2011/07/10/the-games-most-underrated-defense-pairing/
Add to those two: Kulikov who is quickly becoming very good, Campbell, Jovocop and a sheltered Gubranson and you have a very serviceable Dcorps.
Miles ahead of the Oilers for sure.
I see it mentioned daily, maybe we could see a list of all of Snow's "screw ups".
Snow's biggest mistake was the Dipietro contract. Pretty massive mistake.
Past that he's been reasonably solid (although I didn't like Neiderreiter where they picked him, but the jury will be out on that for a few years yet)
Getting a very good young Dman in Wishart for Rollie was an excellent move.
Picking Nabakov off waivers will result in him either playing for NYI or being traded for a good asset.
For what he has to work with, I don't think Snow is terrible.
They just need to cut off Dipietro's leg so he can be LTIR for the rest of his career.
August 16 2011, 08:52AM
Snow's biggest mistake was the Dipietro contract. Pretty massive mistake.
Past that he's been reasonably solid (although I didn't like Neiderreiter where they picked him, but the jury will be out on that for a few years yet)
Getting a very good young Dman in Wishart for Rollie was an excellent move.
Picking Nabakov off waivers will result in him either playing for NYI or being traded for a good asset.
For what he has to work with, I don't think Snow is terrible.
They just need to cut off Dipietro's leg so he can be LTIR for the rest of his career.
Yup, Dipis contract is pretty much all that sticks out to me.... and that really has the looks of an ownership move to me.
I do always get a laugh when Oiler fans take a stab at Snow though.
August 16 2011, 09:00AM
@OB1 Team Yakopov - F.S.T.N.F
Hey, It's not like I've said "Garth Snow sux, Tambellini is WAAAAAAAAAAAAAAAAY better."
They are both GMs of bottom feeder NHL teams. Sorry, but neither deserve praise. Both should be considered subpar.
August 16 2011, 09:10AM
Hey, It's not like I've said "Garth Snow sux, Tambellini is WAAAAAAAAAAAAAAAAY better."
They are both GMs of bottom feeder NHL teams. Sorry, but neither deserve praise. Both should be considered subpar.
Oh ya I agree, it wasn't so much directed at you more so just a generalization of those that comment on Snow.
That said, I actually think he's done a resonably good job.
If Oil stay relatively healthy, then they should be fine.
27th on PP is from a loss of firepower and familiarity. They were 10%! less than Canucks
29th on PK is from lousy goaltending, (can old goalies get across the crease fast enough to stop the back door goal??), poor face off numbers and again, injuries. This is an easier fix. Surprisingly, Chicago and San Jose suck here too.
Unfortunately, THN is probably right. Winning 10 more games than last year is reasonable.
That still puts Edm at last in West, about 25 overall.
I will say this only once. "If" the oilers stay healthy, they will end up 8-10th in the west. If injuries once again plague this team they will end up last or darn close to it.
I am a die hard Oilers fan. The worst team I ever watched play in Edmonton was back in the mid 90's. They were terrible. Glen Sather was about to bolt to New York where he finished trading the 80's Oilers to. The only thing he did before he left worth talking about was he rounded up a bunch of first round picks from the league that were not getting any chances to play. Some of those players developed into nhlers. Today we have a similar type of young Oilers. The difference is we have a bunch of future elite players. The addition of quality veterns to address many team weaknesses from last year. The only weakness I see is in net. Unless Dubnyk gets his chance to be number one. I believe the Oilers will improve much faster because they play the best teams in the league on a regular basis. Playoff bound in 2011-2012.
August 16 2011, 09:40AM
@OB1 Team Yakopov - F.S.T.N.F
I think that much like Tambellini he inherited a team that has been performing worse the longer he stays on.
This is great for the future (as long as you can capitalize on draft position), but I could lead an NHL team into the basement and have enough smarts to realize that I cant mess up drafting high.
I think often people don't appreciate how much things can swing from season to season.
In my opinion it is unfair to look at last year's wins or goal differential and draw a conclusion based on how much they have improved from that marker. What if last year's result was below "what they should have obtained"?
Momentum can be huge in any sport throughout the season.
Take a minute to compare this year's roster to the Oiler's roster in 07-08 and 08-09. There are some differences but I don't see us being that far off of two Oiler teams that were within 3 and 6 points of making the playoffs.
I expect them to finish anywhere between 8th and 14th in the West this year.
I think that much like Tambellini he inherited a team that has been performing worse the longer he stays on.
This is great for the future (as long as you can capitalize on draft position), but I could lead an NHL team into the basement and have enough smarts to realize that I cant mess up drafting high.
This years club miles ahead of last two seasons talent wise , with sophmore jinx unlikely to be a problem . They now have enough of a reputable base from which to build upon . Take a player like even newbie Hunter Tremblay . 2.11 points /game that was/is an all round specialist in University hockey ! 5 shorthanded goals - this player will be hard pressed not to make our team . He fills so many specialist roles at a high performance level. Another newbie i am high on is Musil to make club this season . Oilers can't help but be much better than last season and should be surprise of the season . Book it !! Better odds on Oilers than stock market ? HMMM - early retirement possibilty here ?
You know who never gets the credit for being terrible that he deserves? Bryan Murray. Tambellini and Snow took bad teams and kept them bad, but could at least argue that they laid the foundation for things to get better.
Murray took over Muckler's team right after they made it to the finals; he was handed an excellent team. Surely he was handed some bad breaks (Heatley, A lot of people thought Redden was the keeper of his big 2), but the fact remains that they got worse every year until they finally finished in the least desirable spot in hockey, 6th from the bottom.
Anyone could keep a bad team bad, it's something special to jump off the penthouse and still miss the basement.
August 16 2011, 01:25PM
Tremblay and Musil would need to be SPECTACULAR in order to make this club based on Contract status alone. There arent very many openings.
As it stands Petry might be sent down to OKC because Chorney will have to clear waivers if HE goes. I dont see how Musil will take a spot away from one of Whitney, Smid, Gilbert, Peckham, Barker, Sutton, or Petry. That's 7 defensemen.
And where will Tremblay take his spot from? The only real opening for a new player is at C and the best competition for that will be between Lander/RNH/Brule/VV. 3 have played pro hockey and 1 is this year's reigning number one overall selection.
I dont see any realistic place for either of the guys you mentioned to earn a spot unless they make it impossible for Renney to cut them.
August 16 2011, 02:50PM
Any chance the Oilers could land Ryan Johansen for that one soft spot left at center ice?
Now that Howson has a No.1 center in Carter, could they be tempted to part with Johansen for a player who will make them a better club this coming season, perhaps a right winger? This would give Columbus two pretty good lines going into the coming season.
August 16 2011, 02:54PM
@Quicksilver ballet
I thought that during the Carter deal Howson said that he was constantly asked about Johansen and he refused to give in. I dont think he refused to move him during that deal just so he could turn around and move him for a weaker package later on.
I think it's a pipe dream to land Johansen.
August 16 2011, 02:56PM
Pretty tough to compare Snow and Tambo when Garth's had 2 more years to work with than Tambellini.
If the Oilers are still a last place team 2 years from now, then you could make the comparison, otherwise, that doesn't seem fair.
August 16 2011, 03:05PM
I thought that during the Carter deal Howson said that he was constantly asked about Johansen and he refused to give in. I dont think he refused to move him during that deal just so he could turn around and move him for a weaker package later on.
I think it's a pipe dream to land Johansen.
I think Holmgren had his choice between Columbus' 8th selection and Ryan Johansen in that Jeff Carter deal. Can't help but feel there's something to be had there for the Oilers. Maybe if the Jackets are teetering around that 8th and final playoff spot something may happen before the trade deadline. To get Hemsky along with Carter without giving up a player on your roster is a pretty big improvement over last yrs club.
August 16 2011, 03:18PM
I dont know if it's that hard. Tambellini has been here long enough that the team is showing the effects of his tenure.
He's made plenty of impact on the roster, positive and negative. Surely he can be compared to Snow in respect to being in charge while the teams go into performance tailspins. They can also be compared in how they overvalued questionable goaltenders (Tambellini dodged a DiPietro sized catastrophe at least).
Columbus, Colorado, and Minny are the only three teams that Edmonton has any type of realistic chance of being better than in the Western conference this year, imho.
Columbus got Carter, but their D and G are as suspect as Edmonton's.
Unless Heatley returns to 50 goal form, Minny is in trouble. I think that losing Burns is really going to hurt them, but their goaltending is miles ahead of Edmonton.
Colorado... lots of shake up over the past year, but if Varlamov turns out to be the real deal, they could be real good. They have more upside on their blueline than Edmonton.
Even if Edmonton stays healthy, it's going to take some luck and some bad things to happen to some other teams for them to not finish last.
Playoffs? Not a chance.
The emotional part of me wants to dump on THN, proclaim they don't follow the Western Conference as much as the follow the East, and they don't know what they're talking about. Unfortunately they were correct in predicting in Oil's dead last finish the past two seasons...so the emotional card doesn't hold.
That said, here's why they're wrong:
- Injuries were a big part of last season. Had Whitney, Horcoff, Eberele, Hemsky and Hall remained injured free, who knows.
- The lack of grit which probably lead to some if not all the injuries. Unfortunately Renney didn't trust having McIntyre fill up a roster spot, demoted Huggy Bear, and overvalued Jacques' ability to send a message to teams taking liberities with our stars. The additions of Eager, Hordchuk, and Sutton should address this. At the very least, Renny should have greater confidence in slotting these 3 in every day spots.
- Inability to win faceoffs. With Belanger, we have at least one proven face-off specialist, Horc a moderate face-off specialist, and hopefully one of the kids or Brule can improve.
- Lack of a Ryan Smyth plugger on the PP, and a cannon shot from the point. Well Smytty is back, so that solves half the PP problem.
Is this enough improvement for playoff spot? I would like to hope so, but probably not. But it should ensure us from finishing dead last again.
August 16 2011, 04:28PM
Whatever...they are probaly not far off...should still be fun to watch the kids find their game for another year, finish last and then have Renney fired for no other reason than we finished last again, have another coach come in and mold the winning team. Dave Tippet anyone? He has to eventually get sick of the situation in Phoenix.
August 16 2011, 05:02PM
I dont think for one bit that the Oilers will finish last. Unless the same bunch get injured again but highly unlikely. I really believe the hockey Gods will show us a little mercy this year. With the exception of LA, Vancouver, Detroit, Chicago and maybe San Jose I think the Oil can skate with rest of the pack in the West. The Oilers are not the only team that has holes to fill.
Seeing this subject is still in play, the Bleacher Report also picked the Oilers at the bottom of the heap:
"Edmonton Oilers fans must be getting used to the pain their team puts them through year after year by now, but their hopes of success being not too far away aren't too far-fetched with the stockpile of young hotshots they've got running in their system.
However they're not going to find that success this year.
Taylor Hall and Jordan Eberle both are coming off extremely promising rookie campaigns, as is Magnus Paajarvi, but let's not forget they've both only played one year in the league and will still need to grow and mature a lot before they'll be able to really carry the team.
Their defense also looks to be completely tattered, boasting nothing more than the combination of Cam Barker and Ryan Whitney to lead them (neither of whom would really even be a first-pairing guy on most other teams).
And while Oiler fans will argue that the addition of Ryan Smyth over the summer is a solid boost of veteran leadership to their team, it looks to me like nothing more than the re-acquisition of a washed-up player in the twilight of his career in a desperate grasp at nostalgia for long-lost winning days.
They will be better this season, but the Oilers will still be sitting on their couches watching the playoffs next spring rather than playing in them."
Pretty damning assessment for the coming season, but pretty accurate. Note, they didn't speak to the goaltending problems.
August 16 2011, 06:08PM
I dont think the Smyth bit is very accurate. He would have led the oiler last year and he was LAs PP goal leader. Not washed up yet.
August 16 2011, 06:33PM
Time for a, Should Steve Tambellini be offered an extension poll.
August 16 2011, 09:25PM
A lot of 'ifs' in the reasons I'm reading for the Oil making the dance. Probably just enough of them to keep them out actually...
August 16 2011, 10:42PM
Pretty tough to compare Snow and Tambo when Garth's had 2 more years to work with than Tambellini.
If the Oilers are still a last place team 2 years from now, then you could make the comparison, otherwise, that doesn't seem fair.
Except the NYI aren't in last anymore, they are making a nice progression.


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Seventh Annual Research Day
Theme: A Survival Guide for the Practice of Healthcare in an Era of Change
Rutgers-School of Health Professions (SHP) Research Day recognizes and emphasizes the importance of research and scholarship conducted in the School of Health Professions. The Research and Scholarship Symposium (previously Research Day) is a great networking opportunity for students and faculty members from different departments within the school and to augment faculty-student interactions through research.
Students will present posters supported by their faculty/mentor. Selected students will provide an oral presentation on their research project.
All students presenting a poster will be considered for the Student Scholarship and Research Awards to be presented at Rutgers-SHP's Convocation. Local and distance students are eligible to participate. More information and instructions will be posted below.
When: Monday, May 9, 2016
Where: Piscataway Campus
Time: 10:00am–4:00pm (breakfast will be availabe starting at 9am)
Location: Great Hall and Main Lecture Hall at Robert Wood Johnson Medical School
Research and Scholarship Symposium Abstract Submission
Please note that all posters should have the Rutgers SHP or RBHS logo in the upper left-hand corner (click here for Rugters' logo website/downloads), and should be sized to fit 36 x 48 rectangle (36 inches tall and 48 inches wide). The SAMPLE template provides a traditional format, but please feel free to add, delete or modify the outline provided. Additionally, you do not have to use this format and can take creative liberty in designing your own poster. Printing quality is important and you should have between 150-200 pixels per inch in order to avoid distortion.
Trifold and binder clips will be provided for you at the Research and Scholarship Symposium. You are responsible for the printing of the poster. Set-up for posters begins at 9:00 a.m. Select student teams will be chosen to present to the larger audience. These students will be notified in advance of the Research and Scholarship Symposium.
Information for Judges
Archived Research Day events


text: Best Hi Tea Places in Abbottabad Posted by FCPK Editor on Nov 02, 2013 And as promised here it is: the guide to the best Hi Tea places in Abbottabad, from business or corporate meetings to family get together's, get your money's worth by learning from our experience.Families usually enjoy buffet on sunday and on various occassions. Find the top restaurants and hotels that offer Hi Tea in Abbottabad, Pakistan. Get phone numbers, prices, addresses and timings for the restaurants that serve hi-tea. - Best Restaurants in Abbottabad Serving Buffet Categories Restaurants Launch Coverage Foodie Experiences FCPK In Media Featured Review Interviews Latest In Your City Video Blogs Occasions and Celebrations EatOye in Media


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STARmeterSEE RANK
Down 3,335,669 this week
Michael Gilbert (I) (1912–2006)
Michael Gilbert was born on July 17, 1912 in Billinghay, Lincolnshire, England. He was a writer, known for The Men from Room 13 (1959), Breakout (1959) and Das tödliche Patent (1963). He was married to Roberta Marsden and Roberta Mary Marsden. He died on February 8, 2006 in London, England. See full bio »
Died:(age 93) in London, England, UK


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Mitochondrial targeted antioxidant transgene product MnSOD delivered by plasmid liposomes has shown to be an effective ionizing irradiation damage protector and mitigator but requires hours for in vivo gene product production. A small molecule mitochondrial targeted hemlgramacidin (GS)-nitroxide, JP4-039, is a rapid acting total body irradiation (TBI) damage mitigator, and is an effective topical therapeutic for beta irradiation burns and combined injury (irradiation/bone wound) in C57BL/6HNsd mice. The First Specific Aim tests the hypothesis that enhanced radiation mitigation by mitochondrial targeted GS-nitroxide can be improved by carbon nanocapsule binding and endothelial or epithelial targeted liposome delivery. The Second Specific Aim tests the hypothesis that mitochondrial targeted small molecules (GS-NOS-1) XJB-5-133 and MCF-201- 89 will be effective radiation damage mitigators, also enhanced by carbon nanocapsule, and liposome targeted delivery. The Third Specific Aim tests the hypothesis that the small molecule p53/mdm2/mdm4 inhibitors, BEB55 and BEB59 are effective TBI mitigators, enhanced by carbon nanocapsule, and liposome targeting. Optimized radiation mitigation by each drug in adult male and female, 21 day old prepubescent mice, E13 pregnant females and their progeny will then be combined in a multidrug regimen. Positive controls for acute and late effects will be mini circle MnSOD-plasmid liposomes, and dual transgene MnSOD, mt-CAT-plasmid liposomes. Methods include timed-release degradable carbon nanocapsule bound small molecules, endothelial p-CAM targeted or, epithelial i-CAM, targeted liposomes, organ specific intestinal, marrow, and lung mitochondrial drug uptake quantitation by ESR, phamnacokinetics studies, total body irradiation, and organ histopathology. A long term goal is the development of a practical topical/transdermally delivered mitigator that is effective when given 24 hrs or later after TBI. These studies should establish and optimize novel multidrug mitochondrial targeting for radiation damage mitigation applicable for use in radiation counter terrorism.
We propose to iprove both the pharmacokinetics and pharmacodynamics of GS-nitroxide as a radiation mitigator and determined additive/synergisitc effects for two other novel mitochondrial-targeted small molecule radiation mitigators: [GS (nitric oxide synthetase)-NOS] inhibitors (NOS-1) and p53/mdm2/mdm4 inhibitors through drug delivery via biodegradable carbon nanocapsules in endothelial or epithelial cell targeted liposomes
|Liu, Bing; Bhatt, Divesh; Oltvai, Zoltán N et al. (2014) Significance of p53 dynamics in regulating apoptosis in response to ionizing radiation, and polypharmacological strategies. Sci Rep 4:6245
|Kalash, Ronny; Berhane, Hebist; Yang, Yong et al. (2014) Improved survival of mice after total body irradiation with 10 MV photon, 2400 MU/min SRS beam. In Vivo 28:1-12
|Kagan, Valerian E; Kapralov, Alexandr A; St Croix, Claudette M et al. (2014) Lung macrophages "digest" carbon nanotubes using a superoxide/peroxynitrite oxidative pathway. ACS Nano 8:5610-21
|Kagan, Valerian E; Chu, Charleen T; Tyurina, Yulia Y et al. (2014) Cardiolipin asymmetry, oxidation and signaling. Chem Phys Lipids 179:64-9
|Amoscato, A A; Sparvero, L J; He, R R et al. (2014) Imaging mass spectrometry of diversified cardiolipin molecular species in the brain. Anal Chem 86:6587-95
|Jiang, Jianfei; Bakan, Ahmet; Kapralov, Alexandr A et al. (2014) Designing inhibitors of cytochrome c/cardiolipin peroxidase complexes: mitochondria-targeted imidazole-substituted fatty acids. Free Radic Biol Med 71:221-30
|Stoyanovsky, D A; Sparvero, L J; Amoscato, A A et al. (2014) Improved spatial resolution of matrix-assisted laser desorption/ionization imaging of lipids in the brain by alkylated derivatives of 2,5-dihydroxybenzoic acid. Rapid Commun Mass Spectrom 28:403-12
|Tyurina, Yulia Y; Domingues, Rosario M; Tyurin, Vladimir A et al. (2014) Characterization of cardiolipins and their oxidation products by LC-MS analysis. Chem Phys Lipids 179:3-10
|Chu, Charleen T; Bayýýr, Hulya; Kagan, Valerian E (2014) LC3 binds externalized cardiolipin on injured mitochondria to signal mitophagy in neurons: implications for Parkinson disease. Autophagy 10:376-8
|Tyurina, Yulia Y; Poloyac, Samuel M; Tyurin, Vladimir A et al. (2014) A mitochondrial pathway for biosynthesis of lipid mediators. Nat Chem 6:542-52
Showing the most recent 10 out of 101 publications


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Used as intermediates in pharmaceutical chemical and organic synthesize. It can react with N-benzoyl-glycine in the presence of 4-(dimethylamino)pyridine and acetic anhydride to produce 4-(1-ethoxy-propylidene)-2-phenyl-4H-oxazol-5-one. It is also applied as a starting material in the synthesis of Triethyl orthoacrylate. Triethyl orthopropionate is also commonly used in Claisen rearrangement reactions that yield γ,δ-unsaturated esters.
Kinichi Tadano, et al. Claisen rearrangement of (Z)-3-deoxy-3-C-[(hydroxymethyl)methylene]-1,2:5,6-di-O-isopropylidene-.alpha.-D-ribo-hexofuranose with triethyl orthopropionate.J. Org. Chem.,1989,54(5), 1223-1227.
Edgar Marquez, et al. Mechanisms of the homogeneous, unimolecular gas-phase elimination kinetics of triethyl orthoacetate and triethyl orthopropionate.J. Phys. Org. Chem.,2008,21(7-8), 666-669.
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Flammable liquid and vapour. Causes skin irritation. Causes serious eye irritation. May cause respiratory irritation.
Sicherheitshinweise: P210-P261-P280-P240-P303+P361+P353-P305+P351+P338-P304+P340-P362-P405-P501a
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June 4, 2015 7:04 am
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The Social Security Administration's Ticket To Work program has been around for over a decade with very low participation. Daniel Bertoni is the director of education, Workforce, and Income Security Issues at GAO. He tells In Depth what changes SSA needs to get this program in gear.
Photos of the amazing, moving, important and amusing things happening in the federal community.
A daily update of important moments in the history of the U.S. government.


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The leading market experts have revised a mean price target of $48.47 on Enbridge Inc (NYSE:ENB) stock. This revised target, which is predicted to be touched in a year, is mean of equity calls posted by the analysts. The distinguished research units have predicted earnings of $0.44 a share for the coming quarter, besides for this fiscal is $1.79.
Technical View
The technical analysis of Enbridge Inc Common Stock substantiates the 50-day MA of Enbridge Inc Common Stock is $42.24, and stock is trading $1.46 points away or +3.47% from 50-day moving average of $42.24. It is trading $2.85 or +6.99% away 200-day moving average of $40.85. Enbridge Inc (NYSE:ENB) 52-week high is $45.77 and the 52-week low is $27.43.
If the stock moves $-2.07, it will mark 52-week high. While in event of +59.31% points dive, a 52-week low is fixed.
Enbridge Inc (NYSE:ENB) P/E ratio came in at 39.76. To get additional details about a firm's equity price, buyers can figure out what's accepted as price to earnings ratio. It is a replication of how high-priced or low-priced an equity is depending on existing or projected earnings growth. The totalling implicates dividing equity market value, or the equity last price, by the mean EPS posted in the earlier four quarters. When fiscal earnings forecasts are accessible, the trailing EPS can be traded with estimates for imminent earnings.
An attractive P/E ratio of a firm in one industry might not be valid for P/E ratio of company operating in another sector. Thus, buyers should evaluate the P/E ratios of equities in a similar industry. Likewise, the comprehensive stock market worth is established in the P/E ratio of the prominent S&P index that implies some of the major firms in the market. By gauging a stock's P/E ratio with renowned index, a buyer can know an equity perceived value against the rest of the market.
Enbridge Inc (NYSE:ENB) PEG ratio is at 46.45. The stock closed at $43.70 in last trading session and so current market cap is $40.83B.
1 Chart Pattern Every Investor Should Know
This little-known pattern preceded moves of 578% in ARWR, 562% in LCI, 513% in ICPT, 439% in EGRX, 408% in ADDUS and more...


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Talk:Buck Rogers: A Life in the Future
|WikiProject Novels
|(Rated Stub-class, Mid-importance)
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BetacommandBot (talk) 18:45, 13 February 2008 (UTC)
Misunderstanding of "original research"[edit]
Original research is just that - synthesis of facts by way of comparing with other facts. Stating a plot element in a book falls under the category of "synopsis" and should not be labeled as OR. 126.96.36.199 (talk) 02:47, 2 February 2013 (UTC)


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|21 year old nudist. Absolutely love being naked around/with other people whether it is sexual or not.
|Outgoing nudist couple out for a good time. We are discrete, but we don't really play safe anymore
|63 yr old married bi male nudist .I luv to be nude on my 40 acres in arizona. I enjoy sex outdoors I enjoy men women or couples
|MWC, nudist, enjoy fun times, no means no, no pressure. Would like to find a bi female or bi or sub male or bi couple to join us in fun. Age is just a number, seeking mature fun people. NO means NO. Non smokers or at least smoke outside. Prefer people who will be nude all the time with us.
|********NOT ACTIVELY LOOKING RIGHT NOW,DUE TO THE RECENT PASSING OF MY WIFE, BUT OPEN TO PLAY IF YOU ARE AN INTERESTED COUPLE OR SELECT SINGLE MALE OR FEMALE(PARTICULARLY A SF WHO LIKES BI MEN) WHO CAN HOST CLOSE TO MY LOCATION (WITHIN 60 MILES OR SO), MOSTLY INTERESTED IN FWB FRIENDS FOR NOW, BUT OPEN TO LTR WITH THE RIGHT WOMAN IF THAT DEVELOPS. I WILL BE CHECKING IN TO THIS SITE TO STAY IN TOUCH AND MEET OTHERS WHO SHARE MY INTERESTS AND MIGHT LIKE TO USE ME AS THEIR LIVE SEX TOY SOMETIME, ALTHOUGH I AM NOT ACTIVELY LOOKING, , IF YOU CONTACT ME I WILL RESPOND TO SEE IF WE CAN GET TOGETHER FOR ME TO SERVE YOU. HOPE TO BE BACK PLAYING MORE WIDELY SOON. Leaving pics for you to enjoy ****************************** ********* **** Passively Bi male sub/switch senior nudist boytoy*** **** I have been a home nudist for years, but am looking for ways to practice clothes free living more widely with friends and acquaintance. ***Seeking couples interested in SAFELY using a senior boytoy/live sex toy for their pleasure: - open to m/m bi play str8 play, I'll follow your lead, or Domme female interested in oral, spanking, and strapon dominance. While primarily intereted in couples, I have and will from time to time play with singles of both sexes*** Shaved or closely trimmed, uncut, loves giving oral to both sexes and is good at it. My free time is very limited, and I'll need some lead time to set up any meeting, but that doesn't mean I'm not for real and very interested in being used as a sexual toy and submitting to your desires, as you'll find out when we are able to get together.
|BI sixties couple We are open to most type of sex.both like both oral. Prefer people over 40 y/o Able to meet most days or night, can host He is 6' 210# grey hair and beard, blue eyes She s 5'4" 160# 40c light brown hair grey eyes. We are nudist, love the nude beach. Wish there where more places around to go nude
|I have been a bisexual man now for a few years. I believe that bisexuality is so right in many ways and I would absolutely love to chat with someone about my views. Bisexuality seems so perfect for me. I am also a nudist; I love to be nude (as I am now writing this). That, like bisexuality seems natural. I went to a place recently that I had an opportunity to sit about in the nude around a swimming pool and had a great conversation with another man. It was just two men chatting, no sexual contact, and no suggestion of sexual intercourse of any type between us. I found it relaxing and beneficial mentally and physically. ually, to me, there is a time and place for everything. I would love to find friends and then proceed from there. It would be nice to have a group of friends, both men and women that is uninhibited to show and share their bodies, no matter the physical looks they have. I go for the psyche over the physical and you will need that to have a great friendship. The physical of a relationship is just a bi product of the psyche. Once I get to know someone and we hit it off I can talk your leg off so be warned. I will chat about a variety of things.
|Anybody know of any clothing optional reserves(nudist or naturist colonies) near the cal central coast?
|I personally see nothing wrong with public nudity. I for one hate wearing clothes and there are no nudist colonies in my city. Not to mention my room mate doesn't want to see me in the natural. We do have a nude beach; however, one has to go down a steep dirt path to enter and I have disabilities which prevent me from doing such.
|Couple looking for other bi nudist. Enjoy the nudist life style and would love to have other nudist join us. I am bi active and wife is only bi curious. Would love to make friends as nudist and then look to adding more pleasures. Sorry not into one night stands.
Nude is always better
|I'm a nudist, but I don't have any opportunities to go to nudist camps or anything like that, and I have a roommate that would flip out if I walked around nude. So I can only go nakers when he's on vacation
|Thanks Skip! I've been on their website but have yet to visit. Happy to hear from a friendly nudist. Email me here on BP if you'd be interested in making a new nudist friend.
|I am a fair weather nudist and once I get a family I will certainly steer them that way. I think nudist children learn a very healthy attitude towards body image.
|Curious as to how many either grew up in nudist families or are in nudist families? Also of kids grew up in nudist families, how many continue to be nudists?
|Michael, I just saw your Forum Post about nudists. I am a nudist and live in LA, but fly my plane around So. Cal in search of good nudist places. Avilla Beach comes to mind in your area, I believe... Would love an excuse to fly to SLO again to get naked and whatever....
|Looking for a local nudist female or nudist couple /group to hang out with and enjoy being naked ......or a coed masturbation group within the metroplex.
|As a nudist and a member of a family nudist camp, i see nothing wrong with it. It gives the children the appreciation of what the human body. As they get older then can choose to continue to go or not, the parents i know who have brought their children have expressed this as their policy.
|Hey, my husband and I are new to this and wanted to try out a swingers party or a nudist beach. We are from Sacramento but are willing to travel. Can anyone help us find a party or nudist beach,
would really appreciate it
|I'm very familiar with nudist resorts in paticular and I think you'll find Caliente in Lutz Florida geared to the younger crowd. I think though in general you'll just find a wide variety but at least at a nudist resort you know if the physical part is there. It just takes a while to develop the chemistry. Rarely do you have the physical and relationship traits go together easily.
|Being a nudist, I've heard the stories at my club about Sandy Hook, but haven't gotten there yet. Can you give me an update as to the availability and ability to access the beach? I love being in the water and the pool, but love the ocean the best. What's the procedure for getting on the beach for a day to start, and how do I go from there?
And, you're always welcome to come to the nudist club I frequent. As a couple, I don't think you even need to be an AANR member for the first 3 visits. And I'll spring for your day access fee for your first visit if you would like.
|As open minded as I am. I have to say as a younger couple, we would not want to be meeting at a nudist camp. That's not very "easy going" for a get together of IL folks. Not everyone is that open to being nude when first meeting somebody. Just my two cents. But as for in GENERAL, we'd want IL meet rather than people traveling for a few hour get together. But if people want to travel. More props to them =).
But I think a nudist camp is too extreme for my liking. No offense to anyone else. =) I just don't do nudity infront of strangers.
|Although I was not raised as a nudist, I have been one since I moved out of my very conservative family. I have been to many social nudist gatherings where children are present and have met many adults who were raised as children. They all seem to be well adjusted to me. I kind of worry about a society that seems to be perfectly fine with children playing violent video games or watching movies showing human bodies getting blown apart, but freak out about how much their child will be forever traumatized if he walks in while mommy and daddy are making whoopee.
|I love being naked and would love to go to a nudist resort or beach sometime.
For all of you nudists out there, I have a question.
I have been told that for a true nudist it isn't about sex and, in fact, have been told that many resorts prohibit sex in public. I have also been told that it is impolite to stare at another person or to otherwise exhibit the sex angle.
I want to go to a nude resort near here but fear that, at least until I get more accustomed to being nude and around nudes, I would get an unintentional hard on which, again as I have been told, violates proper etiquette. So, how does one handle that type of situation?
|For Me It Was actuallyThe very Day That I Had Turned !8 Months Old And Had Sucked My Fathers,His Boyfriend's And My Uncles Big Beautiful s And They In Turn ted To Suck On My Then Very Little Dinky They also Did My Balls And They even fucked my ass which has always been something That I have Very Totally d Ever Since, They also let me suck on each and every one of them,They Also Shared Me with very many other bisexual
and gay men and little boy nudists which had been and continues to be A Very special connection to our families bisexual nudist hearitage; Which very,very many bisexual nudist families have shared for very many long centuries plus and very many generations,And even now,I can still feel those moths sucking on my little dinky and balls and knowing for shure right then that this and bisexuality were something that I really was going to persue for the rest of my life,And I Have.
|Nudist Resorts
We are looking for set up summer plans to visit nudist resorts, swing, bi friendly events etc. Any recommendations Thanks Also looking to meet up with couples and single women while traveling.
|Naked Women
Being a nudist myself, I've never quite understood why a woman would want to festoon her body with 'sexy' undergarments - the naked form (in all its different shapes and sizes) is more erotic than anything else . Besides, if those 'sexy' clothes attain their desired effect, they'll be off soon anyway! Seems like a lot of money to spend for not too much wearing time. Victoria's Secret doesn't make much money off of me, I'll tell ya!


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The MatrixCam TM video development kit (VDK) is a smart low power, 1080P camera solution that streams video over Wi-Fi and Ethernet. It has been developed specifically to encourage greater video adoption for IoT products, enabling faster time-to-market for IoT developers. Among its key applications are home automation systems, building access devices, baby monitors, advanced lighting controls and smart appliances.
The MatrixCam VDK offers smart streaming capability, with wake-up initiated by Bluetooth Low Energy (BLE) or through the motion-detecting integrated PIR sensor. Upon wake-up, it sends a push notification to an assigned mobile device (smartphone, tablet, etc.) alerting the user that a live stream or a still image is available for viewing. It supports micro SD card slot, and can connect to cloud services such as Amazon Web Services and Google Cloud Platform, with support for both video-on-demand and live streaming. The VDK can be configured via a simple web-based GUI, or using a dedicated mobile application.
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