Perceptual Learning: Complete Transfer Across Retinal Locations
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Elsevier - Publisher Connector Current Biology Vol 18 No 24 R1134 can drive convergence in avian spectacular plumages through this radiation. Proc. Natl. Acad. Sci. USA 101, 11040–11045. morphology [15,16]. long period of evolutionary isolation. 10. Hackett, S.J., Kimball, R.T., Reddy, S., As suggested by Fleischer and Bowie, R.C., Braun, E.L., Braun, M.J., The Loss of an Ancient Lineage his colleagues [4], the Hawaiian Chojnowski, J.L., Cox, W.A., Han, K.L, Harshman, J., et al. (2008). A phylogenomic The misleading taxonomy caused by honeyeater lineage is best classified study of birds reveals their evolutionary history. this convergent evolution has been into its own new family, the Mohoidae. Science 320, 1763–1768. 11. Fuchs, J., Fjeldsa, J., Bowie, R.C.K., Voelker, G., rectified by the new DNA-based Sadly, this is the only avian family and Pasquet, E. (2006). The African warbler genus analyses, which reveal the surprising known to have gone extinct in Hyliota as a lost lineage in the Oscine songbird uniqueness of these Hawaiian birds. its entirety in the past several tree: Molecular support for an African origin of the Passerida. Mol. Phyl. Evol. 39, 186–197. The last sighting of a Hawaiian centuries. Its demise therefore 12. Cibois, A., and Cracraft, J. (2004). Assessing Chaetoptila occurred in 1859, and three represents the loss of a particularly the passerine ‘‘Tapestry’’: phylogenetic relationships of the Muscicapoidea inferred of the four Moho species were likewise divergent evolutionary lineage from nuclear DNA sequences. Mol. Phyl. Evol. extinct by the mid-1900s. The final [19,20], one that we only now 32, 264–273. surviving species, Moho braccatus, recognize for its true level of 13. Willerslev, E., and Cooper, A. (2005). Ancient DNA. Proc. Royal. Soc. Lond. B 272, 3–16. persisted in the highlands of Kauai into uniqueness. 14. Millar, C.D., Huynen, L., Subramanian, S., the late 1980s, but is now almost Mohandesan, E., David, M., and Lambert, D.M. (2008). New developments in ancient genomics. certainly also extinct. The poignancy of References Trends Ecol. Evol. 23, 386–393. this lineage’s decline is captured in 1. Futuyma, D.J. (2005). Evolution (Sunderland, 15. Beecher, W.J. (1951). Convergence in the recordings of the haunting song of the MA: Sinauer Associates). Coerebidae. Wilson Bull. 63, 274–287. 2. Harmon, L.J., Kolbe, J.J., Cheverud, J.M., and 16. Burns, K.J., Hackett, S.J., and Klein, N.K. (2003). last (and mate-less) male Kauaii ‘O’o, Losos, J.B. (2005). Convergence and the Phylogenetic relationships of Neotropical which can be played online from the multidimensional niche. Evolution 59, 409–421. honeycreepers and the evolution of feeding 3. Wake, D.B. (1991). Homoplasy: the result of morphology. J. Avian Biol. 34, 360–370. 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The 3rd edition (London: Christopher Helm). 9. Barker, F.K., Cibois, A., Schikler, P., Hawaiian honeyeaters evolved their Feinstein, J., and Cracraft, J. (2004). Phylogeny nectar-feeding adaptations and and diversification of the largest avian DOI: 10.1016/j.cub.2008.11.006 Perceptual Learning: Complete location or orientation of the lines between training and test greatly Transfer across Retinal Locations reduces or abolishes the effect of practicing Vernier discriminations or other simple discrimination tasks [3–5]. A newly developed ‘double training’ technique demonstrates that practice- Such specificity is unlikely to be dependent improvement in the discrimination of basic visual features will present with more complex stimuli, transfer to a location that has been trained with a different discrimination. such as faces [6,7], that are not restricted to a single retinal location. Dominic M. Dwyer distinctions between complex Research reported by Xiao et al. [8] in multimodal stimuli that are not this issue of Current Biology questions Gibson [1] defined perceptual possible for novices [2] to the fact whether perceptual learning with learning as ‘‘any relatively permanent that extensive practice with very simple visual stimuli is genuinely and consistent change in the simple discriminations, for example specific to particular retinal locations perception of a stimulus array, between the Vernier offsets of two by demonstrating that, with following practice or experience sets of lines, leads to improved appropriate training methods, with this array’’. This definition performance [3]. improvements in discrimination can encompasses practice-dependent There is much evidence that transfer completely across locations. improvements in performance perceptual learning with simple stimuli The issue of whether or not ranging from the observation that can be very specific to the training perceptual learning is specific to experienced wine-tasters can make situation: changes in the retinal trained locations or stimuli is of Dispatch R1135 critical importance because early A B C D visual cortex contains neurons that are more selective to the position and orientation of stimuli than are neurons further down the visual processing stream. Thus, the apparent specificity + + + + of perceptual learning to particular orientations or locations was considered to be very strong evidence that the neural mechanisms E Test after phase 1 training underpinning the improvements must Test after phase 2 training 40 involve the early visual cortex (for example [9,10]). So consistent have the observations of stimulus 30 specificity been that they have, quite justifiably, been taken as a grounding 20 constraint on both empirical investigations and theoretical accounts of perceptual learning (for 10 example [10,11]). Despite this, it was noted over a decade ago that the 0 specificity in perceptual learning might lie in what is learnt, rather than where Mean % improvement the learning takes place [12], and this –10 possibility has only now been directly investigated [8]. –20 A simplified version of the Ori1 Ori1 Ori2 Ori2 Ori1 Ori2 double-training version of the Vernier Loc1 Loc2 Loc1 Loc2 Loc2 Loc1 discrimination task used by Xiao et al. [8] is shown in Figure 1. Initially Current Biology training was given with lines of one orientation in one location (for example, Figure 1. A simplified version of Experiment 3 from [8]. vertical lines in the upper left visual (A) Phase 1 training with Vernier discriminations using vertical lines in the upper left quadrant; quadrant). In line with previous results, orientation 1-location 1 (Ori1Loc1 in (E), which shows the percent improvement over baseline this produced an improvement in in this condition). (B) The transfer test condition orientation 1-location 2 (Ori1Loc2): Vernier discrimination performance, but this discrimination with the original training stimulus at a new location. (C) Transfer test condition improvement did not transfer to orientation 2-location 1 (Ori2Loc1): Vernier discrimination with a new stimulus at the original location. (D) Phase 2 training with Vernier discriminations using horizontal lines in the horizontal lines tested at the original lower left quadrant (orientation 2-location 2, Ori2Loc2). The transfer tests shown in (B) and location or to the vertical lines tested at (C) were performed after both phase 1 and phase 2 training (left and right sides of (E), respec- a different location such as the lower tively). Note, in the actual experiment the locations and line orientations were counterbal- left visual quadrant (see the leftmost anced. (E) Summary results as mean percentage improvement in discrimination performance three bars in Figure 1E). over baseline. The left side shows performance after phase 1 training and the right side shows Subsequently, training was given performance after phase 2 training. with a second orientation at a second location — for example, horizontal locations when the two discriminations [13,14]), but that the critical site for lines in the lower left visual were trained concurrently and when the process rather than the outcome quadrant — before testing with the different types of discrimination, of perceptual learning must be assignment of line orientation to rather than different orientations of more central or complete transfer training location reversed.