Neural correlates of perceptual learning in a sensory-motor, but not a sensory, cortical area
Open accessAbstract
This study aimed to identify neural mechanisms that underlie perceptual learning in a visual-discrimination task. We trained two monkeys (Macaca mulatta) to determine the direction of visual motion while we recorded from their middle temporal area (MT), which in trained monkeys represents motion information that is used to solve the task, and lateral intraparietal area (LIP), which represents the transformation of motion information into a saccadic choice. During training, improved behavioral sensitivity to weak motion signals was accompanied by changes in motion-driven responses of neurons in LIP, but not in MT. The time course and magnitude of the changes in LIP correlated with the changes in behavioral sensitivity throughout training. Thus, for this task, perceptual learning does not appear to involve improvements in how sensory information is represented in the brain, but rather how the sensory representation is interpreted to form the decision that guides behavior.
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TL;DR
In two monkeys trained on a visual motion direction-discrimination task, improved sensitivity to weak motion was accompanied by changes in motion-driven responses in LIP but not MT. The results suggest perceptual learning involved changes in how sensory evidence is interpreted to form a decision, rather than changes in the sensory representation itself.
Key findings
- Discrimination thresholds improved from ~67% to ~15% coherence for monkey C and from ~68% to ~19% coherence for monkey Z, with time constants of 24.6±0.4 and 58.5±2.5 sessions, respectively.
- MT responses did not change significantly with training: neurometric thresholds were ~4% better during versus before training but not significant (monkey C: 25.9% before vs 21.0% during, p=0.1964; monkey Z: 21.1% before vs 18.4% during, p=0.4762).
- LIP responses developed a coherence × time interaction that grew with training and correlated with behavioural threshold (partial correlation r=−0.42, p<10−5 for monkey C; r=−0.35, p=0.0049 for monkey Z).
- MT choice probability increased from 0.493±0.011 in the first 30% of sessions to 0.549±0.018 in the last 30%, and LIP–behaviour model correlations were r=0.59 (p<10−12) for monkey C and r=0.60 (p<10−9) for monkey Z, whereas MT–behaviour correlations were not significant.
Methods
- Two naïve rhesus monkeys performed a one-interval, two-alternative direction-discrimination task using random-dot motion, reporting direction with an eye movement while MT and LIP neurons were recorded during training.
- Behavioural data were fit with a time-dependent cumulative Weibull function to estimate discrimination threshold; MT and LIP responses were normalized and fit with a linear model containing coherence, viewing time, and coherence × time interaction terms.
- Nested models compared behavioural, MT, and LIP data via a common motion-sensitive scaling term; MT neurometric thresholds and choice probabilities were also computed.
Limitations
- The paper states that changes in other extrastriate visual areas carrying motion signals, such as MST, cannot be ruled out.
- The paper states that the exact role of LIP in task performance throughout training is unknown, and multiple brain areas including superior colliculus and FEF show similar sensory-motor responses.
- The text makes evident that only two monkeys were studied, that stimulus configuration varied slightly across sessions because it depended on recorded neuron properties, and that correlated firing among MT pairs was based on a small number of simultaneously recorded pairs (n=8 before, n=9 during training).
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Claims the paper makes
0 of 57 cited sources checked · 0 supported · 0 flagged
- Findingown data§ Abstract
Behavioural sensitivity to weak motion signals improved continuously during training, long after monkeys had acquired the stimulus/response association.
“Behavioural sensitivity to weak motion signals improved continuously during training, long after monkeys had acquired the stimulus/response association.”
Quote in the paperSubstantial claim - Findingown data§ Abstract
During training, improved behavioural sensitivity to weak motion signals was accompanied by changes in motion-driven responses of neurons in LIP but not MT.
“During training, improved behavioural sensitivity to weak motion signals was accompanied by changes in motion-driven responses of neurons in LIP but not MT.”
- Also stated in: § Abstract · plain
“This improvement in behavioural sensitivity corresponded to changes in the responses of neurons in LIP but not MT during motion viewing.”
Quote in the paperSubstantial claim - Also stated in: § Abstract · plain
- Findingown data§ Abstract
The time course and magnitude of the changes in LIP were correlated with the changes in behavioural sensitivity throughout training.
“The time course and magnitude of the changes in LIP were correlated with the changes in behavioural sensitivity throughout training.”
- Also stated in: § Results · plain
“Moreover, LIP activity was correlated with behavioural threshold across sessions after taking into account the high-coherence errors (Fig. 5, left; partial correlation rth,(k3|λ) = −0.42, H0: r=0 using Fisher’s Z transformation, p<10−5 for monkey C; rth,(k3|λ) =−0.35, p=0.0049 for monkey Z)”
Quote in the paperSubstantial claim - Also stated in: § Results · plain
- Findingown data§ Results
Behavioural performance improved steadily for both monkeys with training.
“Behavioural performance improved steadily for both monkeys with training (Fig. 2).”
Quote in the paperNot a substantial claim - Findingown data§ Results
This lack of long-term changes across sessions occurred despite short-term changes within sessions.
“Interestingly, this lack of long-term changes across sessions occurred despite short-term changes within sessions, as has been reported previously (geometric mean±SEM ratio of psychometric thresholds in the second 200 versus the first 200 trials = 0.92±0.22 in monkey C and 0.84±0.29 in monkey Z; ratio of neurometric thresholds in MT in the second 200 versus the first 200 trials = 0.87±0.23 in monk”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [30] Zohary, E., Celebrini, S., Britten, K.H. & Newsome, W.T. Neuronal plasticity that underlies improvement in perceptual performance. Science 263, 1289–1292 (1994). source
Not checked in the free run.
- Findingown data§ Results
There was a slight strengthening of the relationship between the trial-by-trial variability of MT responses and the monkeys' choices with training.
“There was, however, a slight strengthening of the relationship between the trial-by-trial variability of MT responses and the monkeys’ choices with training.”
Quote in the paperSubstantial claim - Findingown data§ Results
None of the three terms in the linear model of MT responses changed significantly with training.
“Moreover, none of the three terms changed significantly with training (linear regression versus session, computed separately for each term, each monkey and before and during training, H0: slope=0, p>0.05 for all conditions).”
Quote in the paperSubstantial claim - Findingown data§ Results
This specificity for the axis of motion was mirrored in neural activity in LIP but not MT.
“This specificity for the axis of motion was mirrored in neural activity in LIP (Fig. 7b, red symbols; monkey C: r=−0.37, p=0.0004; monkey Z: r=−0.29, p=0.0204) but not MT (Fig. 7b, cyan symbols; monkey C: r=0.09, p=0.5439; monkey Z: r=0.22, p=0.2063)”
Quote in the paperSubstantial claim - Findingown data§ Discussion
Responses of the most sensitive MT neurons became increasingly predictive of the monkeys' choices with training.
“However, responses of the most sensitive MT neurons became increasingly predictive of the monkeys’ choices with training.”
Quote in the paperSubstantial claim - Findingown data§ Results
On trials in which the other target was selected, LIP activity tended to decrease gradually during motion viewing and then remain below baseline levels until after the saccade.
“On trials in which the other target was selected, LIP activity tended to decrease gradually during motion viewing and then remain below baseline levels until after the saccade.”
Quote in the paperSubstantial claim - Findingown data§ Results
Other response properties including the width of directional tuning were unaffected by training.
“Other response properties including the width of directional tuning were unaffected by training (Table S1 and Fig. S3).”
Quote in the paperSubstantial claim - Findingown data§ Results
The coherence × time interaction term in LIP began near zero and progressed steadily to more positive values with training.
“A larger effect was seen in the coherence × time interaction term, which began near zero and thus indicated little or no influence of the motion stimulus on the LIP responses at the beginning of training, and progressed steadily to more positive values.”
Quote in the paperSubstantial claim - Findingown data§ Results
The relationship between choice probability and motion sensitivity in MT changed systematically over the course of training such that more sensitive neurons became more predictive of the monkey's direction decision as training progressed.
“Moreover, the relationship between choice probability and motion sensitivity in MT changed systematically over the course of training such that more sensitive neurons became more predictive of the monkey’s direction decision as training progressed (Fig. 3c; correlation coefficient between choice probability and neurometric thresholds of MT neurons in the 1st 1/3 sessions, r=0.02, H0: r=0, p=0.5600”
Quote in the paperSubstantial claim - Findingown data§ Results
The converse was not true: there was no correlation between high-coherence errors and LIP activity after taking into account the changes in behavioural threshold.
“but the converse was not true, with no correlation between high-coherence errors and LIP activity after taking into account the changes in behavioural threshold (rλ,(k3|th) =0.0460, p=0.6198 for monkey C; rλ,(k3|th)= −0.0590, p=0.6011 for monkey Z).”
Quote in the paperSubstantial claim - Findingown data§ Results
For both monkeys, discrimination performance depended on the familiarity of the axis of motion.
“For both monkeys, discrimination performance depended on the familiarity of the axis of motion (Fig. 7).”
Quote in the paperSubstantial claim - Findingown data§ Discussion
For monkeys trained to discriminate the direction of motion of noisy visual stimuli, improvements in discrimination threshold corresponded to changes in sensory-driven responses in area LIP but not MT.
“Our results show that for monkeys trained to discriminate the direction of motion of noisy visual stimuli, improvements in discrimination threshold (i.e., higher sensitivity to weak motion) corresponded to changes in sensory-driven responses in area LIP but not MT.”
Quote in the paperSubstantial claim - Findingown data§ Results
For both monkeys, the coherence- and time-alone terms in the linear model of LIP responses tended to be near zero throughout training.
“For both monkeys, the coherence- and time-alone terms tended to be near zero throughout training (although the coherence term decreased slightly to small negative values with training in both monkeys, and the time term increased slightly to small positive values with training in monkey C).”
Quote in the paperSubstantial claim - Findingcitation§ Results
MT responses before and during training showed a pattern of activation similar to that reported in trained monkeys.
“As illustrated in Fig. 3a, MT responses before and during training showed a pattern of activation similar to that reported in trained monkeys16: motion in the preferred direction elicited an increase in spike rate and motion in the opposite direction elicited a decrease in spike rate to relatively constant values that depended on motion strength and sustained throughout motion viewing.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [16] Britten, K.H., Shadlen, M.N., Newsome, W.T. & Movshon, J.A. The analysis of visual motion: a comparison of neuronal and psychophysical performance. J. Neurosci. 12, 4745–4765 (1992). source
Not checked in the free run.
- Findingown data§ Results
Other behavioural parameters including the shape and time dependence of the psychometric function did not show similar systematic changes with training.
“Other behavioural parameters including the shape and time dependence of the psychometric function did not show similar systematic changes with training (Table S1).”
Quote in the paperNot a substantial claim - Findingown data§ Results
Choice probability increased as a function of training session.
“Choice probability increased as a function of training session (combined data from both monkeys, mean±SEM choice probability = 0.493±0.011 for the first 30% and 0.549±0.018 for the last 30% of sessions).”
Quote in the paperSubstantial claim - Findingown data§ Results
Performance tended to degrade when an unfamiliar stimulus was used.
“For both monkeys, the value of this quantity was negatively correlated with the absolute z score of motion direction (monkey C: correlation coefficients r=−0.39, H0: r=0 using Fisher’s Z transformation p<10−7; monkey Z: r=−0.49, p<10−9), indicating that performance tended to degrade when an unfamiliar stimulus was used.”
Quote in the paperSubstantial claim - Findingown data§ Results
For both monkeys, only the coherence-alone term in the linear model of MT responses was typically >0.
“For both monkeys, only the coherence-alone term was typically >0, consistent with the idea that MT responses provide evidence about motion direction that depends only on stimulus strength and not viewing time.”
Quote in the paperSubstantial claim - Findingown data§ Results
The monkeys learned to better discriminate weaker and weaker motion signals at shorter viewing times.
“In addition to learning the association, the monkeys learned to better discriminate weaker and weaker motion signals at shorter viewing times.”
Quote in the paperSubstantial claim - Findingown data§ Discussion
In LIP, neurons tuned for saccadic direction were insensitive to visual motion at the beginning of training but developed responses that grew increasingly strongly with motion strength and viewing time as performance improved.
“In LIP, neurons tuned for saccadic direction were insensitive to visual motion at the beginning of training but developed responses that grew increasingly strongly with motion strength and viewing time as performance on the task improved.”
Quote in the paperSubstantial claim - Findingown data§ Results
The neurometric thresholds of neither the whole population nor the 50% most sensitive MT neurons changed systematically as a function of session either before or during training.
“Moreover, the neurometric thresholds of neither the whole population nor the 50% most sensitive MT neurons changed systematically as a function of session either before or during training (linear regression, H0: slope of log threshold versus session=0; before training: monkey C, p=0.7612; monkey Z, p=0.9507, during training: monkey C, whole population, p=0.9498, 50% most sensitive, p=0.9899;”
Quote in the paperSubstantial claim - Findingown data§ Discussion
Motion-driven LIP responses appeared when the monkeys learned the association between motion direction and saccadic response.
“we found that motion-driven LIP responses appeared when the monkeys learned the association between motion direction and saccadic response.”
Quote in the paperSubstantial claim - Findingown data§ Discussion
The study found no evidence for modulation of LIP responses by numerous oculomotor and motivational parameters.
“we found no evidence for such modulation by numerous oculomotor and motivational parameters.”
Quote in the paperSubstantial claim - Interpretationown data§ Abstract
For this task, perceptual learning appears to involve improvements not in how sensory information is represented in the brain but rather how the sensory representation is interpreted to form the decision that guides behaviour.
“Thus, for this task, perceptual learning appears to involve improvements not in how sensory information is represented in the brain but rather how the sensory representation is interpreted to form the decision that guides behaviour.”
- Also stated in: § Abstract · hedged
“The results are consistent with a model in which perceptual improvements result from changes in how sensory evidence is interpreted and used to instruct behaviour, as reflected in LIP, but not in changes to the representation of the evidence itself in MT.”
- Also stated in: § Results · hedged
“Our results suggest that the improvements in behavioural sensitivity result from changes not in the sensory representation but rather its interpretation.”
Quote in the paperSubstantial claim - Also stated in: § Abstract · hedged
- Premisereasoning§ Abstract
MT provides at least some of the motion information used to form the direction decision.
“Thus, MT provides at least some of the motion information used to form the direction decision.”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [16] Britten, K.H., Shadlen, M.N., Newsome, W.T. & Movshon, J.A. The analysis of visual motion: a comparison of neuronal and psychophysical performance. J. Neurosci. 12, 4745–4765 (1992). source
Not checked in the free run.
- cites [17] Britten, K.H., Newsome, W.T., Shadlen, M.N., Celebrini, S. & Movshon, J.A. A relationship between behavioral choice and the visual responses of neurons in macaque MT. Vis. Neurosci. 13, 87–100 (1996). source
Not checked in the free run.
- cites [18] Newsome, W.T. & Pare, E.B. A selective impairment of motion perception following lesions of the middle temporal visual area (MT). J. Neurosci. 8, 2201–2211 (1988). source
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- cites [19] Salzman, C.D., Britten, K.H. & Newsome, W.T. Cortical microstimulation influences perceptual judgments of motion direction. Nature 346, 174–177 (1990). source
Not checked in the free run.
- Premisereasoning§ Abstract
LIP appears to play a role in transforming motion information into a saccadic choice.
“Thus, LIP appears to play a role in transforming motion information into a saccadic choice, although that role might be shared among several oculomotor regions including the frontal eye field and superior colliculus that show similar responses during task performance27, 28.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [24] Shadlen, M.N. & Newsome, W.T. Neural basis of a perceptual decision in the parietal cortex (area LIP) of the rhesus monkey. J. Neurophysiol. 86, 1916–1936 (2001). source
Not checked in the free run.
- cites [25] Roitman, J.D. & Shadlen, M.N. Response of neurons in the lateral intraparietal area during a combined visual discrimination reaction time task. J. Neurosci. 22, 9475–9489 (2002). source
Not checked in the free run.
- cites [26] Hanks, T.D., Ditterich, J. & Shadlen, M.N. Microstimulation of macaque area LIP affects decision-making in a motion discrimination task. Nat. Neurosci. 9, 682–689 (2006). source
Not checked in the free run.
- Premisecitation§ Abstract
Monkeys trained on visual tasks show only moderate or no change in early visual cortex.
“However, monkeys trained on visual tasks show only moderate or no change in early visual cortex7–11.”
Quote in the paperSubstantial claim0 of 5 sources checked- cites [7] Schoups, A., Vogels, R., Qian, N. & Orban, G. Practicing orientation identification improves orientation coding in V1 neurons. Nature 412, 549–553 (2001). source
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- cites [8] Crist, R.E., Li, W. & Gilbert, C.D. Learning to see: experience and attention in primary visual cortex. Nat. Neurosci. 4, 519–525 (2001). source
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- cites [9] Ghose, G.M., Yang, T. & Maunsell, J.H. Physiological correlates of perceptual learning in monkey V1 and V2. J. Neurophysiol. 87, 1867–1888 (2002). source
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- cites [10] Yang, T. & Maunsell, J.H. The effect of perceptual learning on neuronal responses in monkey visual area V4. J. Neurosci. 24, 1617–1626 (2004). source
Not checked in the free run.
- cites [11] Raiguel, S., Vogels, R., Mysore, S.G. & Orban, G.A. Learning to see the difference specifically alters the most informative V4 neurons. J. Neurosci. 26, 6589–6602 (2006). source
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- Premisecitation§ Discussion
Electrophysiological studies in monkeys found little or no changes in V1 with perceptual learning.
“electrophysiological studies in monkeys that found little or no changes in V1 with perceptual learning7–9”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [7] Schoups, A., Vogels, R., Qian, N. & Orban, G. Practicing orientation identification improves orientation coding in V1 neurons. Nature 412, 549–553 (2001). source
Not checked in the free run.
- cites [8] Crist, R.E., Li, W. & Gilbert, C.D. Learning to see: experience and attention in primary visual cortex. Nat. Neurosci. 4, 519–525 (2001). source
Not checked in the free run.
- cites [9] Ghose, G.M., Yang, T. & Maunsell, J.H. Physiological correlates of perceptual learning in monkey V1 and V2. J. Neurophysiol. 87, 1867–1888 (2002). source
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- Interpretationown data§ Results
The changes in sensory-driven LIP responses reflected the improvements in behavioural sensitivity to weak motion signals.
“The changes in sensory-driven LIP responses reflected the improvements in behavioural sensitivity to weak motion signals.”
Quote in the paperSubstantial claim - Premisecitation§ Discussion
Motion sensitivity of MT neurons likely arises from their direct and indirect input from V1.
“the motion sensitivity of MT neurons likely arises from their direct and indirect input from V135”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [35] Movshon, J.A. & Newsome, W.T. Visual response properties of striate cortical neurons projecting to area MT in macaque monkeys. J. Neurosci. 16, 7733–7741 (1996). source
Not checked in the free run.
- Interpretationreasoning§ Discussion
The results appear to rule out learning-induced changes in V1.
“our results appear to rule out learning-induced changes in V1, as well.”
Quote in the paperSubstantial claim - Interpretationown data§ Results
The changes in sensory-driven LIP responses with training did not appear to reflect changes in other motor or motivational parameters known to modulate LIP activity.
“The changes in sensory-driven LIP responses with training also did not appear to reflect changes in other motor (Fig. 5, middle) or motivational parameters (Fig. 5, right) known to modulate LIP activity21, 22.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [21] Platt, M.L. & Glimcher, P.W. Neural correlates of decision variables in parietal cortex. Nature 400, 233–238 (1999). source
Not checked in the free run.
- cites [22] Snyder, L.H., Batista, A.P. & Andersen, R.A. Intention-related activity in the posterior parietal cortex: a review. Vision Res. 40, 1433–1441 (2000). source
Not checked in the free run.
- Interpretationown data§ Results
MT responses provide evidence about motion direction that depends only on stimulus strength and not viewing time.
“For both monkeys, only the coherence-alone term was typically >0, consistent with the idea that MT responses provide evidence about motion direction that depends only on stimulus strength and not viewing time.”
Quote in the paperSubstantial claim - Interpretationcitation§ Discussion
The stimulus specificity of perceptual learning does not necessarily imply changes in primary sensory cortex.
“provides additional evidence that the stimulus specificity of perceptual learning does not necessarily imply changes in primary sensory cortex3, 4.”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationown data§ Results
The systematic change in choice probability is likely to reflect an increasingly selective read-out of activity from MT neurons, particularly those most sensitive to the motion stimulus.
“Thus, the systematic change in choice probability is likely to reflect an increasingly selective read-out of activity from MT neurons, particularly those most sensitive to the motion stimulus.”
Quote in the paperSubstantial claim - Interpretationreasoning§ Discussion
The perceptual improvements corresponded to an increasingly selective read-out of highly sensitive MT neurons by a decision process, represented in LIP, that instructed the behavioural response.
“The results suggest that the perceptual improvements corresponded to an increasingly selective read-out of highly sensitive MT neurons by a decision process, represented in LIP, that instructed the behavioural response.”
Quote in the paperSubstantial claim - Interpretationcitation§ Discussion
The results further support a close relationship between learning and attention.
“The results also further support a close relationship between learning and attention12.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [12] Ahissar, M. & Hochstein, S. Attentional control of early perceptual learning. Proc. Natl. Acad. Sci. USA 90, 5718–5722 (1993). source
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- Premisecitation§ Discussion
Learned sensory-motor associations have been shown to correspond to changes in LIP but not MT.
“Learned sensory-motor associations have been shown to correspond to changes in LIP but not MT41, 42.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [41] Freedman, D.J. & Assad, J.A. Experience-dependent representation of visual categories in parietal cortex. Nature 443, 85–88 (2006). source
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- cites [42] Grunewald, A., Linden, J.F. & Andersen, R.A. Responses to auditory stimuli in macaque lateral intraparietal area. I. Effects of training. J. Neurophysiol. 82, 330–342 (1999). source
Not checked in the free run.
- Interpretationcitation§ Discussion
Decision computations represented in LIP appear to be critical for performance of trained monkeys.
“These decision computations represented in LIP appear to be critical for performance of trained monkeys, an idea supported by a close relationship between LIP responses and reaction times25 and the ability to bias performance using electrical microstimulation of LIP26.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [25] Roitman, J.D. & Shadlen, M.N. Response of neurons in the lateral intraparietal area during a combined visual discrimination reaction time task. J. Neurosci. 22, 9475–9489 (2002). source
Not checked in the free run.
- cites [26] Hanks, T.D., Ditterich, J. & Shadlen, M.N. Microstimulation of macaque area LIP affects decision-making in a motion discrimination task. Nat. Neurosci. 9, 682–689 (2006). source
Not checked in the free run.
- Interpretationown data§ Discussion
The results support the possibility that associative and perceptual learning share common mechanisms.
“These results support the intriguing possibility that associative and perceptual learning might share common mechanisms4, 43.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [4] Dosher, B.A. & Lu, Z.L. Mechanisms of perceptual learning. Vision Res. 39, 3197–3221 (1999). source
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- cites [43] Hall, G. Perceptual and Associative Learning (Oxford University Press, Oxford, 1991). source
Not checked in the free run.
- Rests on an assumption: Learning a stimulus-response association establishes functional connectivity from neurons that represent the sensory stimulus to neurons that control the motor response.
- Study design and conductcitation§ Discussion
The recording and analysis methods could resolve within-session changes in MT sensitivity that were smaller in magnitude than the longer-term changes expected based on behaviour.
“our recording and analysis methods could resolve within-session changes in MT sensitivity30 that were smaller in magnitude than the longer-term changes that we would expect based on behaviour.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [30] Zohary, E., Celebrini, S., Britten, K.H. & Newsome, W.T. Neuronal plasticity that underlies improvement in perceptual performance. Science 263, 1289–1292 (1994). source
Not checked in the free run.
- Interpretationown data§ Discussion
The sensory-driven LIP responses changed in a manner consistent with a decision process that used increasingly sensitive motion evidence to determine the saccadic choice.
“the sensory-driven LIP responses changed in a manner that was consistent with a decision process that used increasingly sensitive motion evidence to determine the saccadic choice.”
Quote in the paperSubstantial claim- The authors state this as an assumption: Sc and Si are normally distributed with variances that scale with their mean.
- The authors state this as an assumption: Sc and Si are independent.
- Premisecitation§ Discussion
Primary somatosensory and auditory cortices in monkeys and humans can show dramatic changes with perceptual learning.
“primary somatosensory and auditory cortices, which in both monkeys and humans can show dramatic changes with perceptual learning5, 6, 37–39.”
Quote in the paperSubstantial claim0 of 5 sources checked- cites [5] Recanzone, G.H., Schreiner, C.E. & Merzenich, M.M. Plasticity in the frequency representation of primary auditory cortex following discrimination training in adult owl monkeys. J. Neurosci. 13, 87–103 (1993). source
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- cites [6] Recanzone, G.H., Merzenich, M.M., Jenkins, W.M., Grajski, K.A. & Dinse, H.R. Topographic reorganization of the hand representation in cortical area 3b owl monkeys trained in a frequency-discrimination task. J. Neurophysiol. 67, 1031–1056 (1992). source
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- cites [37] Pleger, B. et al. Functional imaging of perceptual learning in human primary and secondary somatosensory cortex. Neuron 40, 643–653 (2003). source
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- cites [38] Bao, S., Chang, E.F., Woods, J. & Merzenich, M.M. Temporal plasticity in the primary auditory cortex induced by operant perceptual learning. Nat. Neurosci. 7, 974–981 (2004). source
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- cites [39] Blake, D.T., Heiser, M.A., Caywood, M. & Merzenich, M.M. Experience-dependent adult cortical plasticity requires cognitive association between sensation and reward. Neuron 52, 371–381 (2006). source
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- Interpretationreasoning§ Discussion
The results are consistent with a refinement of direct or indirect ascending projections from MT to LIP.
“Our results are consistent with such a refinement of direct or indirect ascending projections from MT to LIP44, an idea that merits further testing.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [44] Lewis, J.W. & Van Essen, D.C. Corticocortical connections of visual, sensorimotor, and multimodal processing areas in the parietal lobe of the macaque monkey. J. Comp. Neurol. 428, 112–137 (2000). source
Not checked in the free run.
- Rests on an assumption: Learning a stimulus-response association establishes functional connectivity from neurons that represent the sensory stimulus to neurons that control the motor response.
- May combine two claims. 18
- The quote only partly states this claim. 1
- Classified as probably not a substantial claim. 9
May combine two claims.
Findingown data§ ResultsBoth monkeys learned how to perform the task in early sessions and soon could express the visuomotor association for high-coherence stimuli with few or no errors.
“Thus, both monkeys learned how to perform the task in early sessions and soon could express the visuomotor association for high-coherence stimuli with few or no errors.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown data§ ResultsThroughout training, on trials in which the monkey selected the saccade target in the neuron's response field LIP activity tended to increase gradually during motion viewing, stay elevated during the subsequent delay period and then increase just prior to saccade onset.
“Throughout training, on trials in which the monkey selected the saccade target in the neuron’s response field LIP activity tended to increase gradually during motion viewing, stay elevated during the subsequent delay period and then increase just prior to saccade onset.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsSome of these parameters changed with training, but none were consistently correlated with the changes in LIP responsiveness over the course of training for both monkeys.
“Some of these parameters changed with training, but none were consistently correlated with the changes in LIP responsiveness over the course of training for both monkeys (Fig. 5 and Table S1).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsNeurometric thresholds in MT were, on average, ~4% better during versus before training, but the differences were not significant.
“Neurometric thresholds were, on average, ~4% better during versus before training, but the differences were not significant (monkey C: geometric mean threshold coherence before training = 25.9%, during training = 21.0%, Mann-Whitney U test p=0.1964; monkey Z: before = 21.1%; during = 18.4%, p=0.4762).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsThe peri-saccadic activity in LIP tended to be smaller in the first ~15 sessions but then remained relatively stable throughout the rest of training.
“The peri-saccadic activity tended to be smaller in the first ~15 sessions (average activity from −0.5 to −0.1s before saccade onset=18.6 spikes/s from sessions 1–15 and 31.3 spikes/s from sessions 16–160, t-test p=0.0157 for monkey C; 27.0 spikes/s from sessions 1–15 and 48.6 sp/s from sessions 16–130, p=0.0024 for monkey Z) but then remained relatively stable throughout the rest of training (”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsFor both monkeys, the value of the motion-sensitive term grew as a function of training session in a similar manner for data from behaviour and LIP but not MT.
“For both monkeys, the value of this motion-sensitive term grew as a function of training session in a similar manner for data from behaviour and LIP but not MT (Fig. 6b,c; monkey C: correlation coefficient between MT and behaviour r=0.03, H0: r=0 using Fisher’s Z transformation, p=0.7874; LIP and behaviour r=0.59, p<10−12; monkey Z: MT and behaviour r=−0.14, p=0.2911; LIP and behaviour r=0.60, p<”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsThe pattern of coherence-specific changes in LIP was similar to the pattern of coherence-specific behavioural improvements: both the rate of rise of stimulus-driven LIP responses and discrimination performance increased as a function of training session for most non-zero coherences, with greater increases for higher coherences.
“The pattern of coherence-specific changes in LIP was similar to the pattern of coherence-specific behavioural improvements: both the rate of rise of stimulus-driven LIP responses and discrimination performance increased as a function of training session for most non-zero coherences, with greater increases for higher coherences (H0: slope of a linear regression between session number and either”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ DiscussionIn MT, individual neurons responded similarly to motion stimuli before and throughout training, with average sensitivity similar to that of trained monkeys.
“In MT, individual neurons responded similarly to motion stimuli before and throughout training, with, on average, a sensitivity that was similar to that of trained monkeys.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ ResultsThroughout training, performance improved significantly for all non-zero coherences for monkey C and for 12.8-99.9% coherence for monkey Z.
“Indeed, throughout training, performance improved significantly for all non-zero coherences for monkey C and for 12.8–99.9% coherence for monkey Z (linear regression of percent correct per coherence with training session, H0: slope=0, p<0.05).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown data§ ResultsA small number of simultaneously recorded pairs of MT neurons provide preliminary evidence that the degree of correlation is similar before and during training and does not increase over the course of training.
“However, a small number of simultaneously recorded pairs of MT neurons provide preliminary evidence that the degree of correlation is similar before (mean±SEM correlation coefficient r=0.16±0.06, n=8) and during (r=0.18±0.04, n=9) training and does not increase over the course of training (linear regression, H0: slope=0, p=0.5728).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ DiscussionBoth behaviour and LIP responses continued to evolve well after the visuomotor association was established, together reflecting improved sensitivity to weak motion.
“we showed that both behaviour and LIP responses continued to evolve well after the visuomotor association was established, together reflecting improved sensitivity to weak motion.”
Quote in the paperSubstantial claimMay combine two claims. The quote only partly states this claim.
Study design and conductown data§ ResultsWe used a time-dependent cumulative Weibull function to estimate discrimination threshold that takes into account associative (high-coherence) errors and thus can indicate changes in threshold distinct from changes in associative learning or lapses of attention.
“To further quantify improvements in performance with training, we used a time-dependent cumulative Weibull function (Eq. 1 and solid lines in Fig. 2a,b) to estimate the discrimination threshold, which is the motion strength at which the monkey achieved ~82% correct for a 1-s viewing duration, for each session. This function takes into account associative (high-coherence) errors (λ in Eq. 1) and”
Quote in the paperSubstantial claimClassified as probably not a substantial claim. May combine two claims.
Premisecitation§ ResultsIn trained monkeys, MT and LIP play different but complementary roles in forming the direction decision: MT provides sensory evidence used to form the decision, whereas LIP accumulates and interprets the sensory evidence to form the categorical judgment that instructs behaviour.
“In trained monkeys, MT and LIP are thought to play different but complementary roles in forming the direction decision: MT (possibly in tandem with other motion-sensitive areas like the middle superior temporal area, or MST) provides sensory evidence used to form the decision, whereas LIP (possibly in tandem with other sensory-oculomotor areas like the FEF and superior colliculus) accumulates and”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [23] Gold, J.I. & Shadlen, M.N. The neural basis of decision making. Annu. Rev. Neurosci. 30, 535–574 (2007). source
Not checked in the free run.
May combine two claims.
Interpretationown data§ ResultsTraining does not affect the representation of sensory evidence in MT directly but effectively scales its output, providing increasingly sensitive evidence to the decision variable in LIP used to guide behaviour.
“A straightforward interpretation of this result is that training does not affect the representation of sensory evidence in MT directly but effectively scales its output, providing increasingly sensitive evidence to the decision variable in LIP used to guide behaviour.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim.
Study design and conductreasoning§ ResultsWe minimized changes in the stimulus configuration across sessions by selecting neurons with consistent tuning properties, including receptive field location and direction preference of MT neurons and response field location of LIP neurons.
“Within this constraint, we sought to minimize changes in the stimulus configuration across sessions by selecting neurons with consistent tuning properties, including receptive field (RF) location and direction preference of MT neurons and response field location of LIP neurons (Fig. S1).”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationown data§ ResultsPerceptual learning can be specific by virtue of what is learned (in this case, the LIP decision variable) and need not involve changes in the sensory representation (direction-selective responses in area MT).
“which is consistent with the idea that perceptual learning can be specific by virtue of what is learned (in this case, the LIP decision variable) and need not involve changes in the sensory representation (direction-selective responses in area MT)3.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [3] Mollon, J.D. & Danilova, M.V. Three remarks on perceptual learning. Spat. Vis. 10, 51–58 (1996). source
Not checked in the free run.
Classified as probably not a substantial claim.
Premisecitation§ ResultsPerceptual learning is typically specific to the stimulus attributes used during training, including motion direction in a direction-discrimination task.
“Perceptual learning is typically specific to the stimulus attributes used during training, including motion direction in a direction-discrimination task33.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [33] Ball, K. & Sekuler, R. A specific and enduring improvement in visual motion discrimination. Science 218, 697–698 (1982). source
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ ResultsTraining both established and then shaped the sensory-driven responses in LIP sensory-motor neurons.
“Because these neurons were selected based on spatially tuned pre-saccadic activity (see Methods, Fig. S5), this result implies that training both established and then shaped the sensory-driven responses in these sensory-motor neurons.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim. May combine two claims.
Study design and conductreasoning§ ResultsTraining occurred only while the responses of at least one MT or LIP neuron were isolated and recorded, allowing direct, session-by-session comparisons of behavioural and neuronal performance as training progressed.
“Training occurred only while the responses of at least one MT or LIP neuron were isolated and recorded, allowing us to make direct, session-by-session comparisons of behavioural and neuronal performance as training progressed.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ DiscussionLarger effects have been found in V1 of human subjects using fMRI.
“larger effects have been found in V1 of human subjects using fMRI36”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [36] Furmanski, C.S., Schluppeck, D. & Engel, S.A. Learning strengthens the response of primary visual cortex to simple patterns. Curr. Biol. 14, 573–578 (2004). source
Not checked in the free run.
Classified as probably not a substantial claim.
Premisecitation§ DiscussionMT has an established role in task performance.
“the established role of MT in task performance16–19”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [16] Britten, K.H., Shadlen, M.N., Newsome, W.T. & Movshon, J.A. The analysis of visual motion: a comparison of neuronal and psychophysical performance. J. Neurosci. 12, 4745–4765 (1992). source
Not checked in the free run.
- cites [17] Britten, K.H., Newsome, W.T., Shadlen, M.N., Celebrini, S. & Movshon, J.A. A relationship between behavioral choice and the visual responses of neurons in macaque MT. Vis. Neurosci. 13, 87–100 (1996). source
Not checked in the free run.
- cites [18] Newsome, W.T. & Pare, E.B. A selective impairment of motion perception following lesions of the middle temporal visual area (MT). J. Neurosci. 8, 2201–2211 (1988). source
Not checked in the free run.
- cites [19] Salzman, C.D., Britten, K.H. & Newsome, W.T. Cortical microstimulation influences perceptual judgments of motion direction. Nature 346, 174–177 (1990). source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ DiscussionOf the learning-induced changes previously found in V1, attentional modulation appears to play a primary role.
“Of the learning-induced changes that have been found previously in V1, attentional modulation appears to play a primary role8.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [8] Crist, R.E., Li, W. & Gilbert, C.D. Learning to see: experience and attention in primary visual cortex. Nat. Neurosci. 4, 519–525 (2001). source
Not checked in the free run.
May combine two claims.
Interpretationcitation§ DiscussionEarly visual cortical areas are distinguished from primary somatosensory and auditory cortices, which in both monkeys and humans can show dramatic changes with perceptual learning.
“Our results also further distinguish early visual cortical areas from primary somatosensory and auditory cortices, which in both monkeys and humans can show dramatic changes with perceptual learning5, 6, 37–39.”
Quote in the paperSubstantial claim0 of 5 sources checked- cites [5] Recanzone, G.H., Schreiner, C.E. & Merzenich, M.M. Plasticity in the frequency representation of primary auditory cortex following discrimination training in adult owl monkeys. J. Neurosci. 13, 87–103 (1993). source
Not checked in the free run.
- cites [6] Recanzone, G.H., Merzenich, M.M., Jenkins, W.M., Grajski, K.A. & Dinse, H.R. Topographic reorganization of the hand representation in cortical area 3b owl monkeys trained in a frequency-discrimination task. J. Neurophysiol. 67, 1031–1056 (1992). source
Not checked in the free run.
- cites [37] Pleger, B. et al. Functional imaging of perceptual learning in human primary and secondary somatosensory cortex. Neuron 40, 643–653 (2003). source
Not checked in the free run.
- cites [38] Bao, S., Chang, E.F., Woods, J. & Merzenich, M.M. Temporal plasticity in the primary auditory cortex induced by operant perceptual learning. Nat. Neurosci. 7, 974–981 (2004). source
Not checked in the free run.
- cites [39] Blake, D.T., Heiser, M.A., Caywood, M. & Merzenich, M.M. Experience-dependent adult cortical plasticity requires cognitive association between sensation and reward. Neuron 52, 371–381 (2006). source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ DiscussionLIP plays a multitude of cognitive and sensory-motor roles.
“LIP plays a multitude of cognitive and sensory-motor roles that could, in principle, change during training and account for the changes in LIP responses during motion viewing 20–22.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [20] Colby, C.L. & Goldberg, M.E. Space and attention in parietal cortex. Annu. Rev. Neurosci. 22, 319–349 (1999). source
Not checked in the free run.
- cites [21] Platt, M.L. & Glimcher, P.W. Neural correlates of decision variables in parietal cortex. Nature 400, 233–238 (1999). source
Not checked in the free run.
- cites [22] Snyder, L.H., Batista, A.P. & Andersen, R.A. Intention-related activity in the posterior parietal cortex: a review. Vision Res. 40, 1433–1441 (2000). source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ DiscussionIn trained monkeys, multiple brain areas including LIP, the superior colliculus and parts of the prefrontal cortex including the FEF exhibit similar sensory-motor responses during performance of the discrimination task.
“In trained monkeys, multiple brain areas including LIP, the superior colliculus and parts of the prefrontal cortex including the FEF exhibit similar sensory-motor responses during performance of the discrimination task24, 25, 27, 28.”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [24] Shadlen, M.N. & Newsome, W.T. Neural basis of a perceptual decision in the parietal cortex (area LIP) of the rhesus monkey. J. Neurophysiol. 86, 1916–1936 (2001). source
Not checked in the free run.
- cites [25] Roitman, J.D. & Shadlen, M.N. Response of neurons in the lateral intraparietal area during a combined visual discrimination reaction time task. J. Neurosci. 22, 9475–9489 (2002). source
Not checked in the free run.
- cites [27] Horwitz, G.D. & Newsome, W.T. Separate signals for target selection and movement specification in the superior colliculus. Science 284, 1158–1161 (1999). source
Not checked in the free run.
- cites [28] Kim, J.N. & Shadlen, M.N. Neural correlates of a decision in the dorsolateral prefrontal cortex of the macaque. Nat. Neurosci. 2, 176–185 (1999). source
Not checked in the free run.
The paper relies on this without support.
Our understanding of the neural plasticity underlying perceptual learning is incomplete.
“Despite the prevalence of this phenomenon, called perceptual learning, our understanding of the underlying neural plasticity is incomplete.”
The paper relies on this without support.
Changes in higher stages of processing that contribute to decision-making and attention have not been identified directly in the brain.
“However, such changes have not been identified directly in the brain.”
The paper relies on this without support.
Learning a stimulus-response association establishes functional connectivity from neurons that represent the sensory stimulus to neurons that control the motor response.
“Learning a stimulus-response association establishes functional connectivity from neurons that represent the sensory stimulus to neurons that control the motor response.”
The authors state this as an assumption.
Sc and Si are normally distributed with variances that scale with their mean.
“We assume that Sc and Si are normally distributed with variances that scale with their mean”
The authors state this as an assumption.
Sc and Si are independent.
“If Sc and Si are independent and normally distributed”
The authors state this as an assumption.
LIP neurons pool from an unknown number of MT neurons.
“from individual LIP neurons that were assumed to pool from an unknown number of MT neurons”
The authors state this as an assumption.
Behavioural data reflect both MT and LIP contributions.
“from behavioural data assumed to reflect both MT and LIP contributions”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
This accumulation is encoded in the firing rates of LIP neurons with response fields aligned to the choice targets representing the direction alternatives.
“This accumulation is encoded in the firing rates of LIP neurons with response fields aligned to the choice targets representing the direction alternatives (18, 55–57).”
From Representation of Confidence Associated with a Decision by Neurons in the Parietal Cortex (2009) · cites this paper as [57]
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- Authors
- Law, C.-T., & Gold, J. I.
- Affiliations
- University of Pennsylvania (US)
- Corresponding author
- Gold, Joshua I
- ORCID
- Gold, Joshua I
- Venue
- Nature Neuroscience
- Published
- 2008-03-09
- Volume
- 11
- Issue
- 4
- Pages
- 505-513
- ISSN
- 1097-6256 (ISSN-L)
- 1546-1726
- PMC
- PMC2424192
- OpenAlex
- W2103314569
- Semantic Scholar
- 9320464b98ae6d580af8a38d93a505c68c71179d
- Cited by
- 516
- References
- 54
- Citations by year
- 2026: 24 · 2025: 27 · 2024: 25 · 2023: 15 · 2022: 20 · 2021: 20 · 2020: 24 · 2019: 31 · 2018: 29 · 2017: 31 · 2016: 29 · 2015: 36 · 2014: 39 · 2013: 29 · 2012: 43
- FWCI
- 16.24
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- Wellcome Trust
- National Eye Institute (R01EY015260, T32EY007035)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
- Social Sciences › Psychology › Experimental and Cognitive Psychology › Multisensory perception and integration
- Keywords
- perceptual learning, middle temporal area, lateral intraparietal area, saccade selection, motion-sensitive neurons, neuroplasticity, sensorimotor integration, decision-making, macaque
- MeSH terms
- Algorithms, Animals, Brain Mapping, Discrimination Learning, Evoked Potentials, Visual, Macaca mulatta, Motion Perception, Parietal Lobe, Psychomotor Performance, Temporal Lobe
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