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news and views nature neuroscience volume 2 no 8 august 1999 691 with the result that the animal uses the dis- tractors as an (unreliable) reference for judg- ing orientation. It will therefore be impor- tant to determine in future experiments whether the powerful effects reported in this study can be generalized to a wider range of attention-demanding tasks. 1. Distler, C., Boussaoud, D., Desimone, R. & Ungerleider, L. G. J. Comp. Neurol. 334, 125–150 (1993). 2. Felleman, D. J. & Van Essen, D. C. Cereb. Cortex 1, 1–47 (1991). 3. Kaas, J. H. Curr. Biol. 5, 1126–1128 (1995). 4. De Weerd, P., Peralta, M. R., Desimone, R. & Ungerleider, L. G. Nat. Neurosci. 2, 753-758 (1999). 5. Zeki, S. M. Brain Res. 53, 422–427 (1973). 6. Merigan, W. Curr. Biol. 3, 226–229 (1993). 7. Merigan, W. H. Vis. Neurosci. 13, 51–60 (1996). 8. Gallant, J. L., Braun, J. & Van Essen, D. C. Science 259, 100–103 (1993). 9. Desimone, R. & Schein, S. J. Neurophysiol. 57, 835–868 (1987). 10. Walsh, V., Butler, S. R., Carden, D. & Kulikowski, J. J. Behav. Brain Res. 50, 115–126 (1992). 11. Iwai, E. & Mishkin, M. Exp. Neurol. 25, 585–594 (1969). 12. Schiller, P. H. & Lee, K. Science 251, 1251–1253 (1991). 13. Moran, J. & Desimone, R. Science 229, 782–784 (1985). 14. Braun, J. J. Neurosci. 14, 554–567 (1994). 15. Treue, S. & Maunsell, J. H. Nature 382, 539–541 (1996). always appeared in the same location with respect to the fixation point from trial to trial. Consequently, the monkey was not tested for its ability to shift attention from one trial to the next. Second, the distrac- tor stimuli appear in different locations from trial to trial, with some closer to and some farther from fixation than the ori- ented gratings. It will be interesting in future studies to see if this aspect of the task is critical to the observed result. Final- ly, the disruptive effect of the distractors is very marked at low contrasts (<20%) and then does not increase much further at contrasts between 20% and 50% (Fig. 3 of the paper). This can be seen most readily for the more disrupted visual field locations (where V4 and TEO lesions overlap), especially when ignoring the two points at 50% at which monkey M2 could not do the discrimination. In future stud- ies, it will be interesting to determine how the contrast dependence of this effect changes as the distractors are made less or more similar to the test stimuli. The attentional deficits observed in this study are surprisingly profound. One might have anticipated that spatial atten- tion could also be controlled by the dor- sal pathway, whose neurons also show strong attentional modulation 15 , yet these results indicate that the dorsal pathway cannot compensate for the loss of V4 or TEO. This raises the question of whether the role of V4 and TEO in spatial atten- tion is general or task specific. If it turns out that orientation discrimination is per- formed in the ventral pathway (which remains to be shown), the deficits seen here might reflect a task specific role, which could explain why the dorsal path- way cannot provide compensation. If, on the other hand, V4 or TEO lesions also affect tasks that are known to be per- formed in the dorsal pathway (for instance, discrimination of motion direc- tion), this would argue for a more global role in attention, perhaps through top- down influences on earlier cortical areas. There are several further complications to the interpretation of these findings. One possibility, suggested by psychophysical experiments in my own laboratory, is that the loss of sensitivity may reflect a retinotopic but nonspecific increase in neural noise, pre- sumably in lower areas that are closely con- nected to the lesioned area. Another unin- teresting possibility is that the deficits might reflect a disruption in the monkey’s ability to judge orientation relative to nearby visual cues (for example, the edge of the display), Fig. 2. Two important stimulus configurations used by De Weerd and colleagues. (a) Stimulus configuration for orien- tation discrimination with no distractor disks present. (b) Stimulus configuration with dis- tracting disks present. To reproduce the con- ditions under which the monkeys viewed these stimuli, the figure should be held at a distance of 8.5 inches from the eye while fixating the round fixation dot in the upper left of each figure. While viewing each figure this way, the reader should try to determine if the target grating is vertically oriented or is rotated slightly clockwise or coun- terclockwise. Signaling myopia Clear vision requires that the cornea and the lens focus light onto the retina. If the shape of the eye produces a focal plane behind the retina, the eye is myopic, and the image appears blurred. In many developing animals, the sharp- ness of retinal images regulates the growth of the sclera (the outer connec- tive tissue sheath of the eye), so that axial eye length is matched to the refractive power of the lens and cornea. In this issue (pages 706–712), Andy Fischer and colleagues suggest a candi- date for this growth-regulating signal. They found that expression of the immediate-early gene ZENK (red) in a particular class of chick retinal amacrine cells (containing glucagon) was regu- lated by image defocus. ZENK expression was suppressed by several experimental conditions that enhance eye growth (such as a lens that blurs vision), but increased under conditions that suppress it, such as lens removal. This suggests that these amacrine cells could be involved in focus-induced changes in ocular growth and refraction. Kalyani Narasimhan © 1999 Nature America Inc. • http://neurosci.nature.com © 1999 Nature America Inc. • http://neurosci.nature.com

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nature neuroscience • volume 2 no 8 • august 1999 691

with the result that the animal uses the dis-tractors as an (unreliable) reference for judg-ing orientation. It will therefore be impor-tant to determine in future experimentswhether the powerful effects reported in thisstudy can be generalized to a wider rangeof attention-demanding tasks.

1. Distler, C., Boussaoud, D., Desimone, R. &Ungerleider, L. G. J. Comp. Neurol. 334,125–150 (1993).

2. Felleman, D. J. & Van Essen, D. C. Cereb.Cortex 1, 1–47 (1991).

3. Kaas, J. H. Curr. Biol. 5, 1126–1128 (1995).

4. De Weerd, P., Peralta, M. R., Desimone, R. &Ungerleider, L. G. Nat. Neurosci. 2, 753-758(1999).

5. Zeki, S. M. Brain Res. 53, 422–427 (1973).

6. Merigan, W. Curr. Biol. 3, 226–229 (1993).

7. Merigan, W. H. Vis. Neurosci. 13, 51–60(1996).

8. Gallant, J. L., Braun, J. & Van Essen, D. C.Science 259, 100–103 (1993).

9. Desimone, R. & Schein, S. J. Neurophysiol. 57,835–868 (1987).

10. Walsh, V., Butler, S. R., Carden, D. & Kulikowski,J. J. Behav. Brain Res. 50, 115–126 (1992).

11. Iwai, E. & Mishkin, M. Exp. Neurol. 25,585–594 (1969).

12. Schiller, P. H. & Lee, K. Science 251,1251–1253 (1991).

13. Moran, J. & Desimone, R. Science 229,782–784 (1985).

14. Braun, J. J. Neurosci. 14, 554–567 (1994).

15. Treue, S. & Maunsell, J. H. Nature 382,539–541 (1996).

always appeared in the same location withrespect to the fixation point from trial totrial. Consequently, the monkey was nottested for its ability to shift attention fromone trial to the next. Second, the distrac-tor stimuli appear in different locationsfrom trial to trial, with some closer to andsome farther from fixation than the ori-ented gratings. It will be interesting infuture studies to see if this aspect of thetask is critical to the observed result. Final-ly, the disruptive effect of the distractorsis very marked at low contrasts (<20%)and then does not increase much furtherat contrasts between 20% and 50% (Fig.3 of the paper). This can be seen mostreadily for the more disrupted visual fieldlocations (where V4 and TEO lesionsoverlap), especially when ignoring the twopoints at 50% at which monkey M2 couldnot do the discrimination. In future stud-ies, it will be interesting to determine howthe contrast dependence of this effectchanges as the distractors are made less ormore similar to the test stimuli.

The attentional deficits observed inthis study are surprisingly profound. Onemight have anticipated that spatial atten-tion could also be controlled by the dor-sal pathway, whose neurons also showstrong attentional modulation15, yet theseresults indicate that the dorsal pathwaycannot compensate for the loss of V4 orTEO. This raises the question of whetherthe role of V4 and TEO in spatial atten-tion is general or task specific. If it turnsout that orientation discrimination is per-formed in the ventral pathway (whichremains to be shown), the deficits seenhere might reflect a task specific role,which could explain why the dorsal path-way cannot provide compensation. If, onthe other hand, V4 or TEO lesions alsoaffect tasks that are known to be per-formed in the dorsal pathway (for

instance, discrimination of motion direc-tion), this would argue for a more globalrole in attention, perhaps through top-down influences on earlier cortical areas.

There are several further complicationsto the interpretation of these findings. Onepossibility, suggested by psychophysicalexperiments in my own laboratory, is thatthe loss of sensitivity may reflect a retinotopicbut nonspecific increase in neural noise, pre-sumably in lower areas that are closely con-nected to the lesioned area. Another unin-teresting possibility is that the deficits mightreflect a disruption in the monkey’s abilityto judge orientation relative to nearby visualcues (for example, the edge of the display),

Fig. 2. Two importantstimulus configurationsused by De Weerd andcolleagues. (a) Stimulusconfiguration for orien-tation discriminationwith no distractor diskspresent. (b) Stimulusconfiguration with dis-tracting disks present.To reproduce the con-ditions under which themonkeys viewed thesestimuli, the figure should be held at a distance of 8.5 inches from the eye while fixating the roundfixation dot in the upper left of each figure. While viewing each figure this way, the reader shouldtry to determine if the target grating is vertically oriented or is rotated slightly clockwise or coun-terclockwise.

Signaling myopiaClear vision requires that the corneaand the lens focus light onto the retina.If the shape of the eye produces a focalplane behind the retina, the eye ismyopic, and the image appears blurred.In many developing animals, the sharp-ness of retinal images regulates thegrowth of the sclera (the outer connec-tive tissue sheath of the eye), so thataxial eye length is matched to therefractive power of the lens and cornea.In this issue (pages 706–712), AndyFischer and colleagues suggest a candi-date for this growth-regulating signal.They found that expression of theimmediate-early gene ZENK (red) in aparticular class of chick retinal amacrinecells (containing glucagon) was regu-lated by image defocus. ZENK expression was suppressed by several experimental conditionsthat enhance eye growth (such as a lens that blurs vision), but increased under conditions thatsuppress it, such as lens removal. This suggests that these amacrine cells could be involved infocus-induced changes in ocular growth and refraction.

Kalyani Narasimhan

© 1999 Nature America Inc. • http://neurosci.nature.com©

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