So much has been written about free will that it is hard to imagine anything new could be said. T... more So much has been written about free will that it is hard to imagine anything new could be said. The notion that free will is an illusion has achieved such wide acceptance among philosophers and neuroscientists that it seems to have acquired the status of dogma. Nonetheless, research in this area continues and this review offers a new analysis of the design limitations and data interpretations of free-will experiments. This review presents 12 categories of fallacious conclusions that some scholars use to promote the idea that free will is an illusion. The next generation of less ambiguous experiments is proposed.
On the second day of class in the Spring semester, students in both groups were informed of the e... more On the second day of class in the Spring semester, students in both groups were informed of the experi-ment, told that any grade differences would be compen-sated for, and told that they should keep an accurate daily log of the time spent studying the course. Students were ...
This research tested morphine and naloxone effects on evoked EEG and unit activity in 3 opiate-re... more This research tested morphine and naloxone effects on evoked EEG and unit activity in 3 opiate-relevant brain areas in response to electrical stimulation of the olfactory bulb in acute, unanesthesized rat preparations. Stimulation evoked clear EEG responses in the amygdala (Amyg) and sometimes in the other areas (caudate and central grey); morphine (15 mg/kg) depressed the Amyg response in some rats, but enhanced it in others, and naloxone usually reversed both kinds of effect. Stimulation caused excitatory unit impulse reponses in the Amyg, and morphine unexpectedly increased the magnitude of the stimulus-evoked excitation; naloxone reversed this enhancement. In control rats, naloxone often decreased the Amyg evoked response. Stimulus-evoked increases in unit activity in the caudate and central grey, when they did occur, were depressed by morphine, but naxoxone had no consistent reversing effect. Both the EEG and unit data indicate that morphine excites, or disinhibits, certain neurons associated with the olfactory-Amyg pathway. There was also some evidence that this pathway contains endorphinergic elements.
In curarized rats, electrical stimulation (6 Hz pulses in 6-sec trains) was delivered either to t... more In curarized rats, electrical stimulation (6 Hz pulses in 6-sec trains) was delivered either to the dorsal hippocampus (Hip), the caudate nucleus (Caud), or to both. Hip and Caud stimulation evoked activity in the motor cortex and the Hip, and the responses commonly grew larger during the stimulation; to a lesser extent, responses occurred in the brainstem reticulum and cerebellar cortex. Combined Hip and Caud stimulation responses were often greater than the responses to stimulation of either Hip or Caud alone, but less than the sum of responses from each single-site stimulation, suggesting occlusive interference because of shared elements in the projection pathways. There were also some instances suggesting mutual facilitation, where the responses to combined stimulation were greater than the sum of the responses to each single-site stimulation. Both kinds of results seem to indicate that the hippocampus, as well as the caudate, has functional connections with motor control areas of the brain and that both areas may interact in motor control.
The innervation of the eyelids is incompletely understood. This is a particular problem for those... more The innervation of the eyelids is incompletely understood. This is a particular problem for those who wish to develop animal models of eyelid dysfunction in humans. Blepharospasm, for example, is a disease of uncontrolled eyelid spasm that is difficult to manage clinically because the aetiology is not understood. The anatomical literature on eyelid innervation is sparse and even conflicting. We attempted to study eyelid innervation, both sensory and motor, with injection of horseradish peroxidase (HRP) into the superior eyelid, inferior eyelid, and bulbar conjunctiva. We used 13 anesthetized weanling cats. Shape and structure of the facial nucleus varied along its rostrocaudal extent, but there was a clear demarcation of lateral and medial division. HRP-filled facial nucleus cells were ipsilateral to the injection site, and label appeared throughout the rostrocaudal length. All injection sites, including bulbar conjunctiva, labelled facial nucleus neurons located with overlapping di...
Explains how to use journal articles in undergraduate education to teach higher level thinking sk... more Explains how to use journal articles in undergraduate education to teach higher level thinking skills and identifies three components of the teaching approach: (1) small group teams; (2) the analytical process; and (3) the computer conference environment. (YDS)
Equine urine was analysed by capillary gas chromatography. The volatile profiles from oestrous an... more Equine urine was analysed by capillary gas chromatography. The volatile profiles from oestrous and dioestrous samples were compared to establish any qualitative or quantitative difference that may have potential value in olfactory communication. Forty-five different volatile compounds were detected. Of these, 17 major compounds were common to all chromatograms. The chemical profile of oestrous urine was distinguished by the presence of a unique peak that was not present in dioestrous samples. Numerous constituents exhibited endocrine dependence: while the concentrations of seix peaks increased at oestrus, the concentrations of another five peaks decreased at the same time. Since oestrous urine, but not dioestrous urine, has been shown to elicit sexual behaviour in the stallion, the unique peak, together with the peaks that were present in increased concentration at oestrus, may represent important chemical signals that stallions use to detect urinary 'oestrous odours'. Stati...
Journal of the American Veterinary Medical Association, 1959
1. J Am Vet Med Assoc. 1959 Nov 1;135:458-62. A history of anthrax. KLEMM DM, KLEMM WR. PMID: 144... more 1. J Am Vet Med Assoc. 1959 Nov 1;135:458-62. A history of anthrax. KLEMM DM, KLEMM WR. PMID: 14409786 [PubMed - indexed for MEDLINE]. MeSH Terms: Anthrax/history*.
We used a serial order nonparametric analysis technique to analyze trains of neuronal action pote... more We used a serial order nonparametric analysis technique to analyze trains of neuronal action potential intervals in terms of classical information theory. We observed a marked correlation between the information theory descriptor, entropy, and several common measures of variability (median interval length, range of interval length, and skewness of the probability density function). The correlation between variability and entropy was accounted for in the following decreasing order: range of intervals, median interval, and skewness. These data suggest a significant relationship between the signal and its variability and entropy, when entropy is calculated by our serial-order pattern detection method.
Techniques were developed for using the classical information theory descriptor, entropy, to quan... more Techniques were developed for using the classical information theory descriptor, entropy, to quantify the "uncertainty" present in neuronal spike trains. Entropy was calculated on the basis of a method that describes the relative relationships of serially ordered interspike intervals by encoding the intervals as a series of symbols, each of which depicts the relative duration of two adjacent spike intervals. Each symbol, or set of symbols has a specific fractional entropy value, derived from its probability of occurrence; moreover, fractional entropy can describe the relative amount of "information" that is associated with the relative location of a given symbol in a string of symbols. Using spike trains from 12 single neurons in the cerebellar cortex of rats, we determined: (1) the mean and S.D. of information content of each symbol in each specific position in a group of symbols (2-4 symbols/group, based on 3-5 adjacent intervals), (2) the 4-symbol groups which...
Summarized herein is the evidence that supports the hypothesis that neuronal action potentials (s... more Summarized herein is the evidence that supports the hypothesis that neuronal action potentials (spike trains) are coded not only in terms of simple discharge rate but also can be serially coded by certain patterns of spike intervals. Based on the relative interval description method of Sherry and Marczynski (1972), our analyses of single-unit activity from cerebellar cortex neurons of rats seem to support three principal categories of conclusions; (1) Serial dependence of intervals does exist. This has been demonstrated with a variety of conventional statistical tests. These serial dependencies have also been shown to be independent of the (nonsequential) interval distribution variability. (2) Information theory is appropriate for evaluating spike trains. We have developed and tested methods for computing a fractional entropy for a given number of adjacent intervals, for assessing the relative fractional entropy of any one interval in a set of intervals, for computing for a group of...
In order to test the influence of drug-induced state changes on serial ordering of interaction po... more In order to test the influence of drug-induced state changes on serial ordering of interaction potential intervals, we evaluated divergence from statistical independence of specific "clusters" of various numbers of adjacent intervals. Such divergence, which was highly significant statistically, was noted using two different methods, one based on relative interval duration, and the other based on categorizing intervals as to short, medium, or long. Before ethanol, significant divergence was seen for clusters of as many as 5-6 adjacent intervals, and this same degree of serial ordering was seen after ethanol. While the apparent cluster length remained unchanged by ethanol, the specific clusters which contributed most to the statistical significance, were generally not the same after ethanol. This suggests that neurons process intervals as "clusters" of a given length but that specific "clusters" may be important "information carriers" in the ner...
The effect of drug-induced state changes on serial ordering of interaction potential intervals wa... more The effect of drug-induced state changes on serial ordering of interaction potential intervals was evaluated by determining if the intervals were generated by a Markov process, and if they were, determining the Markov level (the degree of serial ordering) of the process before and after the injection of an intoxicating dose of ethanol. When categorizing intervals as short, medium, or long, a Markov process of level 4 (i.e., the process has a "memory" for 4 preceding adjacent intervals) was noted for most neurons; this level of serial ordering remained constant after ethanol injections. Many of the specific clusters that made the major contributions to the statistical significance before ethanol did not contribute after ethanol and a new set of clusters made the major contribution. This suggests that neurons may process intervals as "clusters" and the "cluster-length" may be a basic function of the neuron and not dependent on the general state of the org...
Research communications in chemical pathology and pharmacology, 1976
We tested the question of whether ethanol's known action on the cerebellum is direct or indir... more We tested the question of whether ethanol's known action on the cerebellum is direct or indirect, via neural pathways. The cerebellum of rats was denervated by large electrolytic lesions of the peduncles. Ethanol (1 gm/kg, intraperitoneally injected) generally depressed impulse activity of neuronal populations in the cerebellar cortex, whereas such activity increased in saline-injected controls.
So much has been written about free will that it is hard to imagine anything new could be said. T... more So much has been written about free will that it is hard to imagine anything new could be said. The notion that free will is an illusion has achieved such wide acceptance among philosophers and neuroscientists that it seems to have acquired the status of dogma. Nonetheless, research in this area continues and this review offers a new analysis of the design limitations and data interpretations of free-will experiments. This review presents 12 categories of fallacious conclusions that some scholars use to promote the idea that free will is an illusion. The next generation of less ambiguous experiments is proposed.
On the second day of class in the Spring semester, students in both groups were informed of the e... more On the second day of class in the Spring semester, students in both groups were informed of the experi-ment, told that any grade differences would be compen-sated for, and told that they should keep an accurate daily log of the time spent studying the course. Students were ...
This research tested morphine and naloxone effects on evoked EEG and unit activity in 3 opiate-re... more This research tested morphine and naloxone effects on evoked EEG and unit activity in 3 opiate-relevant brain areas in response to electrical stimulation of the olfactory bulb in acute, unanesthesized rat preparations. Stimulation evoked clear EEG responses in the amygdala (Amyg) and sometimes in the other areas (caudate and central grey); morphine (15 mg/kg) depressed the Amyg response in some rats, but enhanced it in others, and naloxone usually reversed both kinds of effect. Stimulation caused excitatory unit impulse reponses in the Amyg, and morphine unexpectedly increased the magnitude of the stimulus-evoked excitation; naloxone reversed this enhancement. In control rats, naloxone often decreased the Amyg evoked response. Stimulus-evoked increases in unit activity in the caudate and central grey, when they did occur, were depressed by morphine, but naxoxone had no consistent reversing effect. Both the EEG and unit data indicate that morphine excites, or disinhibits, certain neurons associated with the olfactory-Amyg pathway. There was also some evidence that this pathway contains endorphinergic elements.
In curarized rats, electrical stimulation (6 Hz pulses in 6-sec trains) was delivered either to t... more In curarized rats, electrical stimulation (6 Hz pulses in 6-sec trains) was delivered either to the dorsal hippocampus (Hip), the caudate nucleus (Caud), or to both. Hip and Caud stimulation evoked activity in the motor cortex and the Hip, and the responses commonly grew larger during the stimulation; to a lesser extent, responses occurred in the brainstem reticulum and cerebellar cortex. Combined Hip and Caud stimulation responses were often greater than the responses to stimulation of either Hip or Caud alone, but less than the sum of responses from each single-site stimulation, suggesting occlusive interference because of shared elements in the projection pathways. There were also some instances suggesting mutual facilitation, where the responses to combined stimulation were greater than the sum of the responses to each single-site stimulation. Both kinds of results seem to indicate that the hippocampus, as well as the caudate, has functional connections with motor control areas of the brain and that both areas may interact in motor control.
The innervation of the eyelids is incompletely understood. This is a particular problem for those... more The innervation of the eyelids is incompletely understood. This is a particular problem for those who wish to develop animal models of eyelid dysfunction in humans. Blepharospasm, for example, is a disease of uncontrolled eyelid spasm that is difficult to manage clinically because the aetiology is not understood. The anatomical literature on eyelid innervation is sparse and even conflicting. We attempted to study eyelid innervation, both sensory and motor, with injection of horseradish peroxidase (HRP) into the superior eyelid, inferior eyelid, and bulbar conjunctiva. We used 13 anesthetized weanling cats. Shape and structure of the facial nucleus varied along its rostrocaudal extent, but there was a clear demarcation of lateral and medial division. HRP-filled facial nucleus cells were ipsilateral to the injection site, and label appeared throughout the rostrocaudal length. All injection sites, including bulbar conjunctiva, labelled facial nucleus neurons located with overlapping di...
Explains how to use journal articles in undergraduate education to teach higher level thinking sk... more Explains how to use journal articles in undergraduate education to teach higher level thinking skills and identifies three components of the teaching approach: (1) small group teams; (2) the analytical process; and (3) the computer conference environment. (YDS)
Equine urine was analysed by capillary gas chromatography. The volatile profiles from oestrous an... more Equine urine was analysed by capillary gas chromatography. The volatile profiles from oestrous and dioestrous samples were compared to establish any qualitative or quantitative difference that may have potential value in olfactory communication. Forty-five different volatile compounds were detected. Of these, 17 major compounds were common to all chromatograms. The chemical profile of oestrous urine was distinguished by the presence of a unique peak that was not present in dioestrous samples. Numerous constituents exhibited endocrine dependence: while the concentrations of seix peaks increased at oestrus, the concentrations of another five peaks decreased at the same time. Since oestrous urine, but not dioestrous urine, has been shown to elicit sexual behaviour in the stallion, the unique peak, together with the peaks that were present in increased concentration at oestrus, may represent important chemical signals that stallions use to detect urinary 'oestrous odours'. Stati...
Journal of the American Veterinary Medical Association, 1959
1. J Am Vet Med Assoc. 1959 Nov 1;135:458-62. A history of anthrax. KLEMM DM, KLEMM WR. PMID: 144... more 1. J Am Vet Med Assoc. 1959 Nov 1;135:458-62. A history of anthrax. KLEMM DM, KLEMM WR. PMID: 14409786 [PubMed - indexed for MEDLINE]. MeSH Terms: Anthrax/history*.
We used a serial order nonparametric analysis technique to analyze trains of neuronal action pote... more We used a serial order nonparametric analysis technique to analyze trains of neuronal action potential intervals in terms of classical information theory. We observed a marked correlation between the information theory descriptor, entropy, and several common measures of variability (median interval length, range of interval length, and skewness of the probability density function). The correlation between variability and entropy was accounted for in the following decreasing order: range of intervals, median interval, and skewness. These data suggest a significant relationship between the signal and its variability and entropy, when entropy is calculated by our serial-order pattern detection method.
Techniques were developed for using the classical information theory descriptor, entropy, to quan... more Techniques were developed for using the classical information theory descriptor, entropy, to quantify the "uncertainty" present in neuronal spike trains. Entropy was calculated on the basis of a method that describes the relative relationships of serially ordered interspike intervals by encoding the intervals as a series of symbols, each of which depicts the relative duration of two adjacent spike intervals. Each symbol, or set of symbols has a specific fractional entropy value, derived from its probability of occurrence; moreover, fractional entropy can describe the relative amount of "information" that is associated with the relative location of a given symbol in a string of symbols. Using spike trains from 12 single neurons in the cerebellar cortex of rats, we determined: (1) the mean and S.D. of information content of each symbol in each specific position in a group of symbols (2-4 symbols/group, based on 3-5 adjacent intervals), (2) the 4-symbol groups which...
Summarized herein is the evidence that supports the hypothesis that neuronal action potentials (s... more Summarized herein is the evidence that supports the hypothesis that neuronal action potentials (spike trains) are coded not only in terms of simple discharge rate but also can be serially coded by certain patterns of spike intervals. Based on the relative interval description method of Sherry and Marczynski (1972), our analyses of single-unit activity from cerebellar cortex neurons of rats seem to support three principal categories of conclusions; (1) Serial dependence of intervals does exist. This has been demonstrated with a variety of conventional statistical tests. These serial dependencies have also been shown to be independent of the (nonsequential) interval distribution variability. (2) Information theory is appropriate for evaluating spike trains. We have developed and tested methods for computing a fractional entropy for a given number of adjacent intervals, for assessing the relative fractional entropy of any one interval in a set of intervals, for computing for a group of...
In order to test the influence of drug-induced state changes on serial ordering of interaction po... more In order to test the influence of drug-induced state changes on serial ordering of interaction potential intervals, we evaluated divergence from statistical independence of specific "clusters" of various numbers of adjacent intervals. Such divergence, which was highly significant statistically, was noted using two different methods, one based on relative interval duration, and the other based on categorizing intervals as to short, medium, or long. Before ethanol, significant divergence was seen for clusters of as many as 5-6 adjacent intervals, and this same degree of serial ordering was seen after ethanol. While the apparent cluster length remained unchanged by ethanol, the specific clusters which contributed most to the statistical significance, were generally not the same after ethanol. This suggests that neurons process intervals as "clusters" of a given length but that specific "clusters" may be important "information carriers" in the ner...
The effect of drug-induced state changes on serial ordering of interaction potential intervals wa... more The effect of drug-induced state changes on serial ordering of interaction potential intervals was evaluated by determining if the intervals were generated by a Markov process, and if they were, determining the Markov level (the degree of serial ordering) of the process before and after the injection of an intoxicating dose of ethanol. When categorizing intervals as short, medium, or long, a Markov process of level 4 (i.e., the process has a "memory" for 4 preceding adjacent intervals) was noted for most neurons; this level of serial ordering remained constant after ethanol injections. Many of the specific clusters that made the major contributions to the statistical significance before ethanol did not contribute after ethanol and a new set of clusters made the major contribution. This suggests that neurons may process intervals as "clusters" and the "cluster-length" may be a basic function of the neuron and not dependent on the general state of the org...
Research communications in chemical pathology and pharmacology, 1976
We tested the question of whether ethanol's known action on the cerebellum is direct or indir... more We tested the question of whether ethanol's known action on the cerebellum is direct or indirect, via neural pathways. The cerebellum of rats was denervated by large electrolytic lesions of the peduncles. Ethanol (1 gm/kg, intraperitoneally injected) generally depressed impulse activity of neuronal populations in the cerebellar cortex, whereas such activity increased in saline-injected controls.
Neuroscientists increasingly believe that we are making progress in understanding how the brain a... more Neuroscientists increasingly believe that we are making progress in understanding how the brain achieves consciousness. Identifying neural correlates of consciousness (NCC) has been the prevailing experimental paradigm. The NCC have been defined as the minimum neuronal mechanisms jointly sufficient for any one specific conscious percept [1]. There are two possible interpretations of this definition, one involving the specific content of consciousness and the other to the overall state of being conscious. Specific content NCCs of conscious recognition of a face, for example, would be the associated impulse discharge in the face recognition area of the neocortex. The overall consciousness state could involve percepts associated with face recognition, such as situational context, psychological valence, and other related factors. Among the more accepted NCCs for the overall state of being consciousness include all the neural processes associated with being awake, such as an activated EEG, a necessary but insufficient basis for consciousness. We should not confuse wakefulness with consciousness, inasmuch as numerous inattentional blindness studies [2] reveal that we can be awake while being unaware of much in our environment. Being awake, caused by activity of the ascending reticular activating system, only provides the neural state in which neocortically mediated consciousness can emerge. Wakefulness is necessary but not sufficient to assure conscious awareness for any and all attentional targets at any given moment. Consciousness inevitably involves detection and mentally attending of stimulus, memory, or ongoing thought. Research on consciousness has involved great interest in the NCC of long-distance synchrony of gamma frequency oscillations. Another well-studied electrophysiological candidate marker of consciousness is a late (>300 ms after the onset of stimulus), positive, fronto-parietal potential evoked by visual or auditory stimuli, called the P3b [1]. These two most widely used electrophysiological signatures of conscious awareness are closely correlated with selective attention to stimuli, memory, and ongoing thought that is sufficiently novel or relevant to "grab our attention." Here, I wish to suggest neural network scanning as a neglected aspect of consciousness operations. The idea is that the brain may have an ongoing scanning process that enables it to detect novel or relevant cognition as it occurs transiently in real time. When such detection occurs, it likely triggers the coherent synchrony of neural activity that enables conscious awareness. As a starting point, let us consider that the default mode for all thinking is unconscious. A phylogenetic perspective argues for unconsciousness as the default operation of awake brain, inasmuch as lower animals are not likely to have conscious thought. It is clear that much human thinking occurs below the level of conscious awareness, even in sleep where memories of the day's events are being consolidated.
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Papers by W. R. Klemm