Juan Manuel Toro
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Juan Manuel Toro has a total of 52 co-authors. Here are the top co-authors by number of shared publications:
- Núria Sebastián-Gallés
Universitat Pompeu Fabra Barcelona Spain
6 shared publications Marina Nespor
Neuroscience Area, SISSA-International School for Advanced Studies
5 shared publications- Alexandre Celma-Miralles
Center for Music in the Brain, Department of Clinical Medicine, Aarhus University & The Royal Academy of Music Aarhus/Aalborg, Aarhus, Denmark
4 shared publications - Paola Crespo-Bojorque
Center for Brain and Cognition, Universitat Pompeu Fabra
4 shared publications - Antoni Rodríguez-Fornells
Department of Cognition, Development and Educational Psychology, Institute of Neurosciences, University of Barcelona, Barcelona, Spain
3 shared publications Paola Crespo-Bojorque
Universitat Pompeu Fabra, Barcelona, Spain
3 shared publications
Latest Publications
- 5 November 2026
- Published by Elsevier in Trends in Cognitive Sciences
A consonant bias in dogs
- Open Access
- 7 January 2026
- Published by Ubiquity Press, Ltd. in Journal of Cognition
- Vol. 9 (1) , 15
ChatGPT Relies More Heavily on Consonants Than on Vowels to Recognize Words
Humans develop biases during language learning. For example, we rely more heavily on consonants than on vowels to identify words. Advances on artificial intelligence have allowed the development of proficient large language models that sometimes mimic humans’ language use. They do so by tracking regularities in natural language datasets that are used to train them. Here we test the hypothesis that tracking such regularities is enough for the emergence of responses that resemble the consonant bias. We asked ChatGPT which of two nonsense words (one with a vowel and one with a consonant change) was more similar to a target word. We observed that the model uses more the consonants than the vowels to perform similarity judgments across words in the two languages that we tested (English and Spanish).
- 7 January 2026
- Published by Ubiquity Press, Ltd. in Journal of Cognition
- Vol. 9 (1) , 15
Open AccessChatGPT Relies More Heavily on Consonants Than on Vowels to Recognize Words
Humans develop biases during language learning. For example, we rely more heavily on consonants than on vowels to identify words. Advances on artificial intelligence have allowed the development of proficient large language models that sometimes mimic humans’ language use. They do so by tracking regularities in natural language datasets that are used to train them. Here we test the hypothesis that tracking such regularities is enough for the emergence of responses that resemble the consonant bias. We asked ChatGPT which of two nonsense words (one with a vowel and one with a consonant change) was more similar to a target word. We observed that the model uses more the consonants than the vowels to perform similarity judgments across words in the two languages that we tested (English and Spanish).
- 23 December 2025
- Published by Springer Nature in Experimental Brain Research
- Vol. 244 (1) , 18
The role of competing grouping patterns and tonal coherence in neural synchronization to musical meter
Musical meter emerges from the hierarchical organization of beats, which is influenced by both external acoustic cues and internally driven processes. This study examines neural and behavioral responses to auditory sequences when two simultaneous grouping patterns, melodic repetition and intensity accents, are candidates to establish a ternary or quaternary metrical representation. We also investigated how tonal coherence and tempo affect these metrical representations. To do so, we recorded finger tapping responses and electroencephalograms (EEG) from musically-naïve participants while they listened to musical sequences. The frequency-tagging analyses of the EEG indicated that melodic repetition, compared to intensity accents, enhances neural synchronization to metrical groupings. In contrast, disrupting tonal coherence with microtonal intervals reduces neural responses to both grouping patterns. No neural synchronization to any of these patterns was observed during a grouping-continuation imagery task. Behavioral data from the tapping task revealed a strong preference for quaternary over ternary groupings. These findings provide evidence that a repeating melodic pattern can establish neural responses to its periodicity, which likely scaffolds a metrical representation of the rhythmic sequences in the brain through complex interactions between bottom-up and top-down mechanisms in the formation of musical meter.
- 23 December 2025
- Published by Springer Nature in Experimental Brain Research
- Vol. 244 (1) , 18
The role of competing grouping patterns and tonal coherence in neural synchronization to musical meter
Musical meter emerges from the hierarchical organization of beats, which is influenced by both external acoustic cues and internally driven processes. This study examines neural and behavioral responses to auditory sequences when two simultaneous grouping patterns, melodic repetition and intensity accents, are candidates to establish a ternary or quaternary metrical representation. We also investigated how tonal coherence and tempo affect these metrical representations. To do so, we recorded finger tapping responses and electroencephalograms (EEG) from musically-naïve participants while they listened to musical sequences. The frequency-tagging analyses of the EEG indicated that melodic repetition, compared to intensity accents, enhances neural synchronization to metrical groupings. In contrast, disrupting tonal coherence with microtonal intervals reduces neural responses to both grouping patterns. No neural synchronization to any of these patterns was observed during a grouping-continuation imagery task. Behavioral data from the tapping task revealed a strong preference for quaternary over ternary groupings. These findings provide evidence that a repeating melodic pattern can establish neural responses to its periodicity, which likely scaffolds a metrical representation of the rhythmic sequences in the brain through complex interactions between bottom-up and top-down mechanisms in the formation of musical meter.
- 1 January 2025
- Published by Oxford University Press (OUP) in Journal of Language Evolution
- Vol. 10 (1)
A focus on vowels in young human infants and animals: early sensitivities in language processing
Very early in life, from a few weeks to 5 months of age, human infants tend to focus on vowels over consonants to process words. In the present review, we discuss recent evidence suggesting that, just like young infants, nonhuman animals also tend to focus on vowels to identify sequences of sounds. This early use of vowels to recognize words might be linked to acoustic properties that are orthogonal to lexical processing. For example, vowels tend to be more salient than consonants, in the sense that vowels tend to be longer, more stable and are produced with more intensity than consonants. Thus, the recent data with nonhuman animals sheds a new light on this early stage of phoneme processing as the result of biological, and evolutionary relevant predispositions.
- 1 January 2025
- Published by Oxford University Press (OUP) in Journal of Language Evolution
- Vol. 10 (1)
A focus on vowels in young human infants and animals: early sensitivities in language processing
Very early in life, from a few weeks to 5 months of age, human infants tend to focus on vowels over consonants to process words. In the present review, we discuss recent evidence suggesting that, just like young infants, nonhuman animals also tend to focus on vowels to identify sequences of sounds. This early use of vowels to recognize words might be linked to acoustic properties that are orthogonal to lexical processing. For example, vowels tend to be more salient than consonants, in the sense that vowels tend to be longer, more stable and are produced with more intensity than consonants. Thus, the recent data with nonhuman animals sheds a new light on this early stage of phoneme processing as the result of biological, and evolutionary relevant predispositions.
- Open Access
- 29 May 2024
- Published by The Royal Society in Royal Society Open Science
- Vol. 11 (5) , 221181
Do rats ( Rattus norvegicus ) perceive octave equivalence, a critical human cross-cultural aspect of pitch perception?
Octave equivalence describes the perception that two notes separated by a doubling in frequency have a similar quality. In humans, octave equivalence is important to both music and language learning and is found cross-culturally. Cross-species studies comparing human and non-human animals can help illuminate the necessary pre-conditions to developing octave equivalence. Here, we tested whether rats ( Rattus norvegicus ) perceive octave equivalence using a standardized cross-species paradigm. This allowed us to disentangle concurring hypotheses regarding the evolutionary roots of this phenomenon. One hypothesis is that octave equivalence is directly connected to vocal learning, but this hypothesis is only partially supported by data. According to another hypothesis, the harmonic structure of mammalian vocalizations may be more important. If rats perceive octave equivalence, this would support the importance of vocal harmonic structure. If rats do not perceive octave equivalence, this would suggest that octave equivalence evolved independently in several mammalian clades due to a more complex interplay of different factors such as—but not exclusively—the ability to vocally learn. Evidence from our study suggests that rats do perceive octave equivalence, thereby suggesting that the harmonic vocal structure found in mammals may be a key pre-requisite for octave equivalence. Stage 1 approved protocol: the study reported here was originally accepted as a Registered Report and the study design was approved in Stage 1. We hereby confirm that the completed experiment(s) have been executed and analysed in the manner originally approved with any unforeseen changes in those approved methods and analyses clearly noted. The approved Stage 1 protocol can be found at: https://osf.io/gvf7c/?view_only=76dc1840f31c4f9ab59eb93cbadb98b7 .
- 29 May 2024
- Published by The Royal Society in Royal Society Open Science
- Vol. 11 (5) , 221181
Open AccessDo rats ( Rattus norvegicus ) perceive octave equivalence, a critical human cross-cultural aspect of pitch perception?
Octave equivalence describes the perception that two notes separated by a doubling in frequency have a similar quality. In humans, octave equivalence is important to both music and language learning and is found cross-culturally. Cross-species studies comparing human and non-human animals can help illuminate the necessary pre-conditions to developing octave equivalence. Here, we tested whether rats ( Rattus norvegicus ) perceive octave equivalence using a standardized cross-species paradigm. This allowed us to disentangle concurring hypotheses regarding the evolutionary roots of this phenomenon. One hypothesis is that octave equivalence is directly connected to vocal learning, but this hypothesis is only partially supported by data. According to another hypothesis, the harmonic structure of mammalian vocalizations may be more important. If rats perceive octave equivalence, this would support the importance of vocal harmonic structure. If rats do not perceive octave equivalence, this would suggest that octave equivalence evolved independently in several mammalian clades due to a more complex interplay of different factors such as—but not exclusively—the ability to vocally learn. Evidence from our study suggests that rats do perceive octave equivalence, thereby suggesting that the harmonic vocal structure found in mammals may be a key pre-requisite for octave equivalence. Stage 1 approved protocol: the study reported here was originally accepted as a Registered Report and the study design was approved in Stage 1. We hereby confirm that the completed experiment(s) have been executed and analysed in the manner originally approved with any unforeseen changes in those approved methods and analyses clearly noted. The approved Stage 1 protocol can be found at: https://osf.io/gvf7c/?view_only=76dc1840f31c4f9ab59eb93cbadb98b7 .
- Open Access
- 2 March 2024
- Published by Springer Nature in Animal Cognition
- Vol. 27 (1) , 1-9
Recognizing structure in novel tunes: differences between human and rats
A central feature in music is the hierarchical organization of its components. Musical pieces are not a simple concatenation of chords, but are characterized by rhythmic and harmonic structures. Here, we explore if sensitivity to music structure might emerge in the absence of any experience with musical stimuli. For this, we tested if rats detect the difference between structured and unstructured musical excerpts and compared their performance with that of humans. Structured melodies were excerpts of Mozart's sonatas. Unstructured melodies were created by the recombination of fragments of different sonatas. We trained listeners (both human participants and Long-Evans rats) with a set of structured and unstructured excerpts, and tested them with completely novel excerpts they had not heard before. After hundreds of training trials, rats were able to tell apart novel structured from unstructured melodies. Human listeners required only a few trials to reach better performance than rats. Interestingly, such performance was increased in humans when tonality changes were included, while it decreased to chance in rats. Our results suggest that, with enough training, rats might learn to discriminate acoustic differences differentiating hierarchical music structures from unstructured excerpts. More importantly, the results point toward species-specific adaptations on how tonality is processed.
- 2 March 2024
- Published by Springer Nature in Animal Cognition
- Vol. 27 (1) , 1-9
Open AccessRecognizing structure in novel tunes: differences between human and rats
A central feature in music is the hierarchical organization of its components. Musical pieces are not a simple concatenation of chords, but are characterized by rhythmic and harmonic structures. Here, we explore if sensitivity to music structure might emerge in the absence of any experience with musical stimuli. For this, we tested if rats detect the difference between structured and unstructured musical excerpts and compared their performance with that of humans. Structured melodies were excerpts of Mozart's sonatas. Unstructured melodies were created by the recombination of fragments of different sonatas. We trained listeners (both human participants and Long-Evans rats) with a set of structured and unstructured excerpts, and tested them with completely novel excerpts they had not heard before. After hundreds of training trials, rats were able to tell apart novel structured from unstructured melodies. Human listeners required only a few trials to reach better performance than rats. Interestingly, such performance was increased in humans when tonality changes were included, while it decreased to chance in rats. Our results suggest that, with enough training, rats might learn to discriminate acoustic differences differentiating hierarchical music structures from unstructured excerpts. More importantly, the results point toward species-specific adaptations on how tonality is processed.
- 1 January 2024
- Published by Comparative Cognition Society in Comparative Cognition & Behavior Reviews
- Vol. 19, 91-95
Insights from Animals to Build Better Artificial Language Learners
- 1 January 2024
- Published by Comparative Cognition Society in Comparative Cognition & Behavior Reviews
- Vol. 19, 91-95
Insights from Animals to Build Better Artificial Language Learners
- Open Access
- 9 October 2023
- Published by Springer Nature in Scientific Reports
- Vol. 13 (1) , 1-8
Sensitivity to the sonority sequencing principle in rats (Rattus norvegicus)
Albeit diverse, human languages exhibit universal structures. A salient example is the syllable, an important structure of language acquisition. The structure of syllables is determined by the Sonority Sequencing Principle (SSP), a linguistic constraint according to which phoneme intensity must increase at onset, reaching a peak at nucleus (vowel), and decline at offset. Such structure generates an intensity pattern with an arch shape. In humans, sensitivity to restrictions imposed by the SSP on syllables appears at birth, raising questions about its emergence. We investigated the biological mechanisms at the foundations of the SSP, testing a nonhuman, non-vocal-learner species with the same language materials used with humans. Rats discriminated well-structured syllables (e.g., pras) from ill-structured ones (e.g., lbug) after being familiarized with syllabic structures conforming to the SSP. In contrast, we did not observe evidence that rats familiarized with syllables that violate such constraint discriminated at test. This research provides the first evidence of sensitivity to the SSP in a nonhuman species, which likely stems from evolutionary-ancient cross-species biological predispositions for natural acoustic patterns. Humans’ early sensitivity to the SSP possibly emerges from general auditory processing that favors sounds depicting an arch-shaped envelope, common amongst animal vocalizations. Ancient sensory mechanisms, responsible for processing vocalizations in the wild, would constitute an entry-gate for human language acquisition.
- 9 October 2023
- Published by Springer Nature in Scientific Reports
- Vol. 13 (1) , 1-8
Open AccessSensitivity to the sonority sequencing principle in rats (Rattus norvegicus)
Albeit diverse, human languages exhibit universal structures. A salient example is the syllable, an important structure of language acquisition. The structure of syllables is determined by the Sonority Sequencing Principle (SSP), a linguistic constraint according to which phoneme intensity must increase at onset, reaching a peak at nucleus (vowel), and decline at offset. Such structure generates an intensity pattern with an arch shape. In humans, sensitivity to restrictions imposed by the SSP on syllables appears at birth, raising questions about its emergence. We investigated the biological mechanisms at the foundations of the SSP, testing a nonhuman, non-vocal-learner species with the same language materials used with humans. Rats discriminated well-structured syllables (e.g., pras) from ill-structured ones (e.g., lbug) after being familiarized with syllabic structures conforming to the SSP. In contrast, we did not observe evidence that rats familiarized with syllables that violate such constraint discriminated at test. This research provides the first evidence of sensitivity to the SSP in a nonhuman species, which likely stems from evolutionary-ancient cross-species biological predispositions for natural acoustic patterns. Humans’ early sensitivity to the SSP possibly emerges from general auditory processing that favors sounds depicting an arch-shaped envelope, common amongst animal vocalizations. Ancient sensory mechanisms, responsible for processing vocalizations in the wild, would constitute an entry-gate for human language acquisition.
- Open Access
- 9 February 2022
- Published by Springer Nature in Animal Cognition
- Vol. 25 (4) , 951-960
Detecting surface changes in a familiar tune: exploring pitch, tempo and timbre
Humans recognize a melody independently of whether it is played on a piano or a violin, faster or slower, or at higher or lower frequencies. Much of the way in which we engage with music relies in our ability to normalize across these surface changes. Despite the uniqueness of our music faculty, there is the possibility that key aspects in music processing emerge from general sensitivities already present in other species. Here we explore whether other animals react to surface changes in a tune. We familiarized the animals (Long–Evans rats) with the “Happy Birthday” tune on a piano. We then presented novel test items that included changes in pitch (higher and lower octave transpositions), tempo (double and half the speed) and timbre (violin and piccolo). While the rats responded differently to the familiar and the novel version of the tune when it was played on novel instruments, they did not respond differently to the original song and its novel versions that included octave transpositions and changes in tempo.
- 9 February 2022
- Published by Springer Nature in Animal Cognition
- Vol. 25 (4) , 951-960
Open AccessDetecting surface changes in a familiar tune: exploring pitch, tempo and timbre
Humans recognize a melody independently of whether it is played on a piano or a violin, faster or slower, or at higher or lower frequencies. Much of the way in which we engage with music relies in our ability to normalize across these surface changes. Despite the uniqueness of our music faculty, there is the possibility that key aspects in music processing emerge from general sensitivities already present in other species. Here we explore whether other animals react to surface changes in a tune. We familiarized the animals (Long–Evans rats) with the “Happy Birthday” tune on a piano. We then presented novel test items that included changes in pitch (higher and lower octave transpositions), tempo (double and half the speed) and timbre (violin and piccolo). While the rats responded differently to the familiar and the novel version of the tune when it was played on novel instruments, they did not respond differently to the original song and its novel versions that included octave transpositions and changes in tempo.