Cited by 463Animal Vocal Communication and BehaviorNeuroscience and Music PerceptionLanguage and cultural evolution

Convergent transcriptional specializations in the brains of humans and song-learning birds

Open access
Pfenning, Andreas R., Hara, Erina, Whitney, Osceola, Rivas, Miriam V., Wang, Rui, Roulhac, Petra L., Howard, Jason T., Wirthlin, Morgan +17 more·2014·Science

Abstract

Song-learning birds and humans share independently evolved similarities in brain pathways for vocal learning that are essential for song and speech and are not found in most other species. Comparisons of brain transcriptomes of song-learning birds and humans relative to vocal nonlearners identified convergent gene expression specializations in specific song and speech brain regions of avian vocal learners and humans. The strongest shared profiles relate bird motor and striatal song-learning nuclei, respectively, with human laryngeal motor cortex and parts of the striatum that control speech production and learning. Most of the associated genes function in motor control and brain connectivity. Thus, convergent behavior and neural connectivity for a complex trait are associated with convergent specialized expression of multiple genes.

AI snapshot

Written from the paper’s full text.

TL;DR

The paper compares brain transcriptomes of vocal-learning birds and humans and identifies convergent gene expression specializations in song and speech brain regions, especially songbird RA with human laryngeal motor cortex and Area X with human putamen/caudate. It argues that convergent vocal-learning behavior and neural connectivity are accompanied by convergent specialized expression of multiple genes.

Key findings

  • Dynamic programming alignment of anatomical gene expression specialization trees found songbird Area X best aligned to human putamen, ventral striatum to human nucleus accumbens, and RA to the human central sulcus within primary motor cortex (PrG_cs); HVC and LMAN aligned weakly to cingulate cortex, with their highest hypothesized matches to Wernicke’s and Broca’s areas but without significant permutation/FWER support.
  • Statistical tests confirmed Area X–putamen and RA–PrG_cs similarities; 288 genes contributed to shared avian–human striatum specialization, 342 to avian pallium–human cortex, 78 to Area X–putamen, 40 to RA–PrG_cs, and 55 mostly nonoverlapping genes (only 3 overlapping with the 40) to RA analogs–human LMC/dLSC.
  • Across all three vocal-learning avian lineages (songbird, parrot, hummingbird), RA analogs showed stronger convergent specialization with human LMC/dLSC than vocal-nonlearning dove/quail arcopallium; 55 genes were identified, with ~91% decreased expression in RA analogs and human LMC/dLSC, and in situ hybridization of 15 genes gave ~95% true positive rate for differential expression in zebra finch RA.
  • Proteomics identified 81 proteins differentially expressed in zebra finch RA; mRNA–protein correlation was modest but significant (R² = 0.18, P = 9.4 × 10⁻⁵), and top disease categories included movement disorders (P = 8.7 × 10⁻¹⁴, 29 genes).

Methods

  • Six experiments were analyzed: laser-microdissected song nuclei from zebra finches; microarray samples of Area X and ventral striatum; RA analogs and adjacent motor arcopallium across vocal-learning and vocal-nonlearning birds; brainstem vocal and neck motor neurons; 3702 human samples from 231 subregions of six Allen Human Brain Atlas subjects; and 258 macaque cortical-layer samples from 12 regions of four rhesus macaques.
  • Avian samples used a custom zebra finch Agilent oligoarray v2.2; human samples used a human Agilent 8×60K array containing 7473 human–finch orthologous genes; cross-species avian filtering yielded 3044 genes for multi-lineage comparisons, and macaque comparisons used 4702 orthologs.
  • Analyses included anatomical gene expression specialization trees, pairwise distances and Pearson correlations, a dynamic programming alignment preserving hierarchical structure, permutation/FWER tests, linear regression to count shared specialized genes, functional enrichment, in situ hybridization, and nanoscale capillary tandem mass spectrometry.

Limitations

  • The paper states that correlations were low even between biological replicates (mean zebra finch RA vs. adjacent mAC replicate correlation = 0.22), that the analyses do not show the specializations are functionally related to projections or vocal learning and could be a consequence rather than a cause, that HVC/LMAN convergent specializations were not significant, that human premortem vocalizing state was unknown, that brainstem vocal/neck motor neurons could not be aligned to proposed human homologs because they were not isolated in the Allen data, and that macaque data were limited in regions, cortical layers, and ortholog number.
  • Evident from the text: cross-species microarray orthology and filtering may miss genes; human samples were collected 10 to 30 hours postmortem, which may affect RNA profiles; avian samples were from silent adult males, limiting behavioral-state and sex coverage; and no causal manipulation tests gene function in vocal learning.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1126/science.1256846

Cites

116 of 116 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 97 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    The mean correlation between biological replicates of zebra finch RA relative to the adjacent mAC specialization is 0.22.

    “For example, the mean correlation between biological replicates of zebra finch RA relative to the adjacent mAC specialization is 0.22.”
    Quote in the paperNot a substantial claim
  • Findingown data

    The dynamic programming algorithm found an optimal alignment between the brain gene expression specialization trees of songbird and human.

    “The dynamic programming algorithm found an optimal alignment between the brain gene expression specialization trees of songbird (Fig. 2A) and human (Fig. 2B; see also Table 1).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Findingown data

    Songbird striatal regions (Area X and VS) were confirmed by both methods to have the most similar specialized gene expression to the human striatum.

    “In particular, songbird striatal regions (Area X and VS) were confirmed by both methods to have the most similar specialized gene expression to the human striatum (Fig. 3A and fig. S3A; Table 1 and table S3A).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Findingown data

    The mAC lateral to RA, which does not participate in song production and does not make direct projections, aligned weakly to human superior rostral gyrus of frontal lobe.

    “The mAC lateral to RA, which does not participate in song production (9, 51) and does not make direct projections (52, 53), aligned weakly to the human superior rostral gyrus (SRoG) of the frontal lobe (Fig. 2B).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Findingown data

    Pearson correlations between specializations were low, even in biological replicates, because of noise and large numbers of genes that are not specialized in expression.

    “We found that the Pearson correlations between specializations were low, even in biological replicates, because of noise and large numbers of genes that are not specialized in expression.”
    Quote in the paperSubstantial claim
  • May combine two claims. 39
  • Classified as probably not a substantial claim. 14
  • The quote only partly states this claim. 2
  • May combine two claims.

    Findingown data

    Avian pallial regions were confirmed to be most similar to human cortex, with lower significance by permutation analyses (P = 0.068) and FWER analyses (0.218) but with the highest number of specialized genes in common of all regions compared.

    “The avian pallial regions were confirmed to be most similar to the human cortex, with a lower significance by the permutation analyses (P = 0.068) and corresponding FWER analyses (0.218) but with the highest number of specialized genes in common of all regions compared (Fig. 3B and fig. S3B; Table 1 and table S3B).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Both HVC and LMAN aligned best to the parietal part of the cingulate cortex, but the correlations were low.

    “Both HVC and LMAN aligned best to the parietal part of the cingulate cortex, but the correlations were low (Fig. 2B).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    At high levels of the alignment, songbird telencephalic regions corresponded most closely to human telencephalon and songbird brainstem nuclei corresponded to human brainstem nuclei.

    “At high levels of the alignment, the songbird telencephalic regions corresponded most closely to the human telencephalon (Tel) and the songbird brainstem nuclei corresponded to human brainstem nuclei.”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Within the telencephalon, songbird pallial regions corresponded most closely to human cortex, and songbird striatal regions corresponded to human striatum.

    “Within the telencephalon, the songbird pallial regions corresponded most closely to the human cortex (Cx), and the songbird striatal regions corresponded to the human striatum.”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    The songbird pallial regions sampled did not have a strong similarity to human claustrum and had an inverse gene expression relationship with the amygdala.

    “The songbird pallial regions sampled did not have a strong similarity to the human claustrum (Cl) and had an inverse gene expression relationship with the amygdala (Amg), meaning that the specialization was in the opposite direction (Table 1).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Of the regions hypothesized to be similar to HVC or LMAN, the highest correlation values were HVC with Wernicke's auditory speech processing region and LMAN with Broca's speech production and processing region.

    “None of the regions hypothesized to be similar to HVC or LMAN had a strong secondary match (correlations > 0.09), but of the hypothesized regions, the highest correlation values were HVC with Wernicke’s auditory speech processing region and LMAN with Broca’s speech production and processing region (Table 1).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    For HVC and LMAN, of the hypothesized human brain regions, the highest similarities were to Wernicke's area (plus Heschl's and transverse auditory gyri) and Broca's area, respectively, but the relationships were not significant according to permutation and FWER tests.

    “For HVC and LMAN, of the hypothesized human brain regions, again the highest similarities were to Wernicke’s area (plus Heschl’s and transverse auditory gyri; HG, TG) and Broca’s area, respectively, but the relationships were not significant according to the permutation and FWER tests (Table 1, fig. S3F, and table S3F).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    The top enrichment categories for avian pallium-human cortex shared genes were behavior (64 genes) and neurotransmission (38 genes).

    “The top enrichment categories were behavior (64 genes) and neurotransmission (38 genes; table S5B).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    The RA analogs of all three vocal learners had many more genes specialized in common with the human LMC/dLSC than did the arcopallium of vocal nonlearners, and the number of specialized genes in common was significant only for the vocal learners.

    “Likewise, the RA analogs of all three vocal learners had many more genes specialized in common with the human LMC/dLSC than did the arcopallium of vocal nonlearners, and the number of specialized genes in common was significant only for the vocal learners (Fig. 3, E and F, and table S4, B and C).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    The common specialized genes between all arcopallium samples of each avian species and the entire human frontal lobe were similar across all avian species.

    “For a control analysis, we identified the common specialized genes between all the arcopallium samples of each avian species and the entire human frontal lobe, and found them to be similar across all avian species (Fig. 3G).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    There was a weak, nonsignificant difference in the correlation between the arcopallium regions of avian vocal learners versus nonlearners and macaque layer 5 cells, but a stronger significant correlation between the RA analogs of avian vocal learners versus nonlearners and human LMC/dLSC versus macaque layer 5 cells.

    “Comparing the arcopallium samples from each of the avian species with the specializations of human LMC/dLSC versus macaque layer 5, we found a weak, nonsignificant difference in the correlation between the arcopallium regions of avian vocal learners versus nonlearners and macaque layer 5 cells (table S4D), but a stronger significant correlation between the RA analogs of avian vocal learners versus”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    The correlations increased for the vocal-learning avian species, but not for the vocal-nonlearning avian species, when comparisons were made to specific human putative LMC (dLMC and vLMC) and dLSC samples within PrG_cs and PoG_cs relative to the cortex.

    “The correlations increased for the vocal-learning avian species, but not for the vocal-nonlearning avian species, when comparisons were made to the specific human putative LMC (dLMC and vLMC) and dLSC samples within the PrG_cs and PoG_cs relative to the cortex (fig. S4C, triangles and squares); the correlations were still present when the comparison was relative to only the remaining PrG and PoG (”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Both tests confirmed Area X to have the strongest similarity with the putamen but a significant secondary relationship with the adjacent body of the caudate.

    “Among the striatal regions, both tests confirmed Area X to have the strongest similarity with the putamen (Fig. 3C and fig. S3C; Table 1 and table S3C) but a significant secondary relationship with the adjacent body of the caudate, consistent with the hypothesis that Area X is similar to adjacent parts of the caudate and putamen (41).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Both tests confirmed RA to have the strongest similarity to the central sulcus part of human primary motor cortex (PrG_cs) but also to the adjacent primary somatosensory part of the central sulcus (PoG_cs) and to a region where the ventral LMC is located (PrG_sl).

    “Among the pallial regions, both tests confirmed RA to have the strongest similarity to the central sulcus part of the human primary motor cortex (PrG_cs) but also to the adjacent primary somatosensory part of the central sulcus (PoG_cs) and to a region where the ventral LMC is located (PrG_sl) (Fig. 3D and fig. S3D; Table 1 and table S3D).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    LMAN had the best match to layer 4 of the PoG and other cortical regions, contradicting previous hypotheses but consistent with a recent study comparing chicken nidopallium to mouse cortical layers.

    “However, LMAN, also part of the nidopallium, had the best match to layer 4 of the PoG and other cortical regions, contradicting previous hypotheses (41, 67) (Table 1, fig. S5, E and F, and table S3I) but consistent with a recent study comparing chicken nidopallium to mouse cortical layers (68).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • The quote only partly states this claim.

    Findingown data

    The adjacent dLSC showed striking down-regulation of SLIT1 throughout the region.

    “The adjacent dLSC showed striking down-regulation throughout the region (Fig. 7).”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    The anatomical profiles of the genes fell into three categories: expression specialization in the RA analog and human dLMC/dLSC cortex; expression specialization in the RA plus immediately adjacent intermediate arcopallium and LMC plus the adjacent human primary motor cortex; and a more varied pattern of expression within the avian or human brain, but with local differences in avian RA and human

    “The anatomical profiles of the genes fell into three categories (Fig. 6 and figs. S9 and S10): (i) those with expression specialization in the RA analog and human dLMC/dLSC cortex; (ii) those with expression specialization in the RA plus immediately adjacent intermediate arcopallium and LMC plus the adjacent human primary motor cortex; and (iii) those with a more varied pattern of expression”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    The top enrichment categories among the 40 genes shared with motor cortex cells were movement disorders and abnormal morphology of neurons, whereas the 55 genes shared more specifically with LMC/dLSC were related to cognitive behaviors and development of neural connections for different gene subsets.

    “The top enrichment categories among the 40 genes shared with motor cortex cells were movement disorders and abnormal morphology of neurons (table S5D), whereas the 55 genes shared more specifically with LMC/dLSC were related to cognitive behaviors (cognition, learning, and memory) and development of neural connections (neuritogenesis, axon and dendrite growth) for different gene subsets (5 to 15”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims. The quote only partly states this claim.

    Findingown data

    In contrast to Area X and the analogous putamen/caudate regions, where more than 70% of the genes had increased expression, the majority of RA/LMC/dLSC genes had decreased expression.

    “This is in contrast to Area X and the analogous putamen/caudate regions, where more than 70% of the genes had increased expression (fig. S7).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    Although there was a possible stronger relationship of RA with the speech-dominant left LMC/dLSC hemisphere, the right hemisphere was also correlated.

    “On the basis of these results, we analyzed a subset of 10 of the 15 genes by in situ hybridization in our predicted dLMC/dLSC region of the other hemisphere (right) of two human brain specimens used for the microarrays; although we noted a possible stronger relationship of RA with the speech-dominant left LMC/dLSC hemisphere, the right hemisphere was also correlated (SM8).”
    Quote in the paperNot a substantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Findingown data

    40 genes contributed to the shared specialized expression between songbird RA and human PrG_cs; a subset of these (10 genes) contributed ~50% to the shared specialization between songbird RA and macaque layer 5 of motor cortex.

    “We identified 40 genes that contributed to the shared specialized expression between songbird RA and human PrG_cs (table S3D); a subset of these (10 genes) contributed ~50% to the shared specialization between songbird RA and macaque layer 5 of motor cortex (table S3D).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingcitation

    FOXP2 was up-regulated in layer 5 cells of primary motor cortex (M1) in mouse.

    “FOXP2 was also in the opposite direction, with up-regulation in layer 5 cells of primary motor cortex (M1; fig. S14) (73).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [73] Hisaoka T Nakamura Y Senba E Morikawa Y The forkhead transcription factors, Foxp1 and Foxp2, identify different subpopulations of projection neurons in the mouse cerebral cortex Neuroscience 166 551 563 2010 10.1016/j.neuroscience.2009.12.055 20040367

      Not checked in the free run.

  • May combine two claims.

    Premiseown data

    Song-learning birds and humans share independently evolved similarities in brain pathways for vocal learning that are essential for song and speech and are not found in most other species.

    “Song-learning birds and humans share independently evolved similarities in brain pathways for vocal learning that are essential for song and speech and are not found in most other species.”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [1] Petkov CI Jarvis ED Birds, primates, and spoken language origins: Behavioral phenotypes and neurobiological substrates Front Evol Neurosci 4 12 2012 10.3389/fnevo.2012.00012 22912615 PMC3419981

      Not checked in the free run.

    • cites [4] Doupe AJ Kuhl PK Birdsong and human speech: Common themes and mechanisms Annu Rev Neurosci 22 567 631 1999 10.1146/annurev.neuro.22.1.567 10202549

      Not checked in the free run.

    • cites [5] Bolhuis JJ Okanoya K Scharff C Twitter evolution: Converging mechanisms in birdsong and human speech Nat Rev Neurosci 11 747 759 2010 10.1038/nrn2931 20959859

      Not checked in the free run.

    • cites [6] Arriaga G Jarvis ED Mouse vocal communication system: Are ultrasounds learned or innate? Brain Lang 124 96 116 2013 10.1016/j.bandl.2012.10.002 23295209 PMC3886250

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Vocal-learning species share babbling, deafness-induced deterioration of learned vocalizations, dialects, and forebrain circuits that control production and learning of vocalizations.

    “Vocal-learning species also share the presence of babbling, deafness-induced deterioration of learned vocalizations, dialects, and forebrain circuits that control production and learning of vocalizations (1, 4–6).”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [1] Petkov CI Jarvis ED Birds, primates, and spoken language origins: Behavioral phenotypes and neurobiological substrates Front Evol Neurosci 4 12 2012 10.3389/fnevo.2012.00012 22912615 PMC3419981

      Not checked in the free run.

    • cites [4] Doupe AJ Kuhl PK Birdsong and human speech: Common themes and mechanisms Annu Rev Neurosci 22 567 631 1999 10.1146/annurev.neuro.22.1.567 10202549

      Not checked in the free run.

    • cites [5] Bolhuis JJ Okanoya K Scharff C Twitter evolution: Converging mechanisms in birdsong and human speech Nat Rev Neurosci 11 747 759 2010 10.1038/nrn2931 20959859

      Not checked in the free run.

    • cites [6] Arriaga G Jarvis ED Mouse vocal communication system: Are ultrasounds learned or innate? Brain Lang 124 96 116 2013 10.1016/j.bandl.2012.10.002 23295209 PMC3886250

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S1 (asserted).

    Premisecitation

    For vocal learning, it has been hypothesized that superregions of a motor learning or auditory perception pathway specialized into subregions that control production of learned vocalizations.

    “For vocal learning, it has been hypothesized that the superregions of a motor learning or auditory perception pathway have specialized into subregions that control the production of learned vocalizations (9–11).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites [9] Feenders G Molecular mapping of movement-associated areas in the avian brain: A motor theory for vocal learning origin PLOS ONE 3 e1768 2008 10.1371/journal.pone.0001768 18335043 PMC2258151

      Not checked in the free run.

    • cites [10] Margoliash D Distributed representation in the song system of oscines: Evolutionary implications and functional consequences Brain Behav Evol 44 247 264 1994 10.1159/000113580 7842284

      Not checked in the free run.

    • cites [11] Mello CV Vates GE Okuhata S Nottebohm F Descending auditory pathways in the adult male zebra finch (Taeniopygia guttata) J Comp Neurol 395 137 160 1998 10.1002/(SICI)1096-9861(19980601)395:2<137::AID-CNE1>3.0.CO;2-3 9603369

      Not checked in the free run.

  • May combine two claims.

    Premisenone given

    The vocal-learning circuits include a corticostriatal loop and a unique direct connection from motor cortical areas (human LMC and songbird RA) to brainstem vocal motor and respiratory neurons for phonation.

    “These circuits include a corticostriatal loop and a unique direct connection from motor cortical areas [human laryngeal motor cortex (LMC) and songbird robust nucleus of the arcopallium (RA)] to brainstem vocal motor and respiratory neurons for phonation (Fig. 1).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Fig. 1

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    FOXP2 transcription factor is required for vocal learning in both humans and songbirds, suggesting neural circuits for vocal learning across clades may have evolved overlapping molecular mechanisms.

    “The FOXP2 transcription factor is required for vocal learning in both humans (20) and songbirds (21–24), which suggests that neural circuits for vocal learning across clades may have evolved overlapping molecular mechanisms.”
    Quote in the paperSubstantial claim0 of 5 sources checked
    • cites [20] Lai CS Fisher SE Hurst JA Vargha-Khadem F Monaco AP A forkhead-domain gene is mutated in a severe speech and language disorder Nature 413 519 523 2001 10.1038/35097076 11586359

      Not checked in the free run.

    • cites [21] Haesler S FoxP2 expression in avian vocal learners and non-learners J Neurosci 24 3164 3175 2004 10.1523/JNEUROSCI.4369-03.2004 15056696 PMC6730012

      Not checked in the free run.

    • cites [22] Haesler S Incomplete and inaccurate vocal imitation after knockdown of FoxP2 in songbird basal ganglia nucleus Area X PLOS Biol 5 e321 2007 10.1371/journal.pbio.0050321 18052609 PMC2100148

      Not checked in the free run.

    • cites [23] Scharff C Petri J Evo-devo, deep homology and FoxP2: Implications for the evolution of speech and language Philos Trans R Soc London Ser B 366 2124 2140 2011 10.1098/rstb.2011.0001 21690130 PMC3130369

      Not checked in the free run.

    • cites [24] Teramitsu I Kudo LC London SE Geschwind DH White SA Parallel FoxP1 and FoxP2 expression in songbird and human brain predicts functional interaction J Neurosci 24 3152 3163 2004 10.1523/JNEUROSCI.5589-03.2004 15056695 PMC6730014

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Some glutamate receptors have specialized expression in forebrain song and speech areas in birds and humans, but a direct comparison between species was not made.

    “Some glutamate receptors were found to have specialized expression in forebrain song and speech areas in birds (19) and humans (18), but a direct comparison between species was not made.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [18] Amunts K Broca’s region: Novel organizational principles and multiple receptor mapping PLOS Biol 8 e1000489 2010 10.1371/journal.pbio.1000489 20877713 PMC2943440 source

      Not checked in the free run.

    • cites [19] Wada K Sakaguchi H Jarvis ED Hagiwara M Differential expression of glutamate receptors in avian neural pathways for learned vocalization J Comp Neurol 476 44 64 2004 10.1002/cne.20201 15236466 PMC2517240

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    A potential mechanism for convergent evolution of brain systems is that a larger brain structure (superregion) specializes into multiple subregions, each performing a distinct function.

    “A potential mechanism for convergent evolution of brain systems is one in which a larger brain structure (superregion) specializes into multiple subregions, each of which performs a distinct function (7–9).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [7] Oakley TH Rivera AS Genomics and the evolutionary origins of nervous system complexity Curr Opin Genet Dev 18 479 492 2008 10.1016/j.gde.2008.12.002 19152785

      Not checked in the free run.

    • cites [8] Barrett HC A hierarchical model of the evolution of human brain specializations Proc Natl Acad Sci USA 109 suppl 1 10733 10740 2012 10.1073/pnas.1201898109 22723350 PMC3386868

      Not checked in the free run.

    • cites [9] Feenders G Molecular mapping of movement-associated areas in the avian brain: A motor theory for vocal learning origin PLOS ONE 3 e1768 2008 10.1371/journal.pone.0001768 18335043 PMC2258151

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    For echolocating mammals, genome-scale convergence in amino acid sequence was found in many genes involved in hearing.

    “In addition, for echolocating mammals, genome-scale convergence in amino acid sequence was recently found in many genes involved in hearing (31).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [31] Parker J Genome-wide signatures of convergent evolution in echolocating mammals Nature 502 228 231 2013 10.1038/nature12511 24005325 PMC3836225

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    The authors did not find a strong correlation between avian striatal regions (Area X and VS) and human pallidum, supporting the conclusion that the avian regions are predominantly striatal.

    “Although some avian striatal regions (Area X and VS) are known to have a sparse population of pallidum-like neurons (46–48), we did not find a strong correlation with the human pallidum (Table 1), supporting the conclusion that the avian regions are predominantly striatal (33, 44).”
    Quote in the paperSubstantial claim0 of 5 sources checked
    • cites [46] Carrillo GD Doupe AJ Is the songbird Area X striatal, pallidal, or both? An anatomical study J Comp Neurol 473 415 437 2004 10.1002/cne.20099 15116398

      Not checked in the free run.

    • cites [47] Reiner A Laverghetta AV Meade CA Cuthbertson SL Bottjer SW An immunohistochemical and pathway tracing study of the striatopallidal organization of area X in the male zebra finch J Comp Neurol 469 239 261 2004 10.1002/cne.11012 14694537

      Not checked in the free run.

    • cites [48] Farries MA Perkel DJ A telencephalic nucleus essential for song learning contains neurons with physiological characteristics of both striatum and globus pallidus J Neurosci 22 3776 3787 2002 11978853 PMC6758399

      Not checked in the free run.

    • cites [33] Zhang G Comparative genomics reveals insights into avian genome evolution and adaptation Science 346 1311 1320 2014 10.1126/science.1251385 25504712 PMC4390078

      Not checked in the free run.

    • cites [44] Reiner A The Avian Brain Nomenclature Forum: Terminology for a new century in comparative neuroanatomy J Comp Neurol 473 E1 E6 2004 19626136 PMC2713747

      Not checked in the free run.

    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationcitation

    The lower significance for avian pallium-human cortex similarity could possibly reflect greater differences between avian pallium and human cortex than between avian and human striatum.

    “The lower significance could possibly reflect greater differences between the avian pallium and human cortex than between the avian and human striatum (33, 47).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [33] Zhang G Comparative genomics reveals insights into avian genome evolution and adaptation Science 346 1311 1320 2014 10.1126/science.1251385 25504712 PMC4390078

      Not checked in the free run.

    • cites [47] Reiner A Laverghetta AV Meade CA Cuthbertson SL Bottjer SW An immunohistochemical and pathway tracing study of the striatopallidal organization of area X in the male zebra finch J Comp Neurol 469 239 261 2004 10.1002/cne.11012 14694537

      Not checked in the free run.

    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Interpretationown data

    These results support the cortex hypothesis and contradict the claustrum-amygdala hypothesis of homologies between avian and mammalian pallial brain regions.

    “These results support the cortex hypothesis (33, 43, 44) and contradict the claustrum-amygdala hypothesis of homologies between avian and mammalian pallial brain regions (33, 45).”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [33] Zhang G Comparative genomics reveals insights into avian genome evolution and adaptation Science 346 1311 1320 2014 10.1126/science.1251385 25504712 PMC4390078

      Not checked in the free run.

    • cites [43] Jarvis ED Global view of the functional molecular organization of the avian cerebrum: Mirror images and functional columns J Comp Neurol 521 3614 3665 2013 23818122 PMC4145244

      Not checked in the free run.

    • cites [44] Reiner A The Avian Brain Nomenclature Forum: Terminology for a new century in comparative neuroanatomy J Comp Neurol 473 E1 E6 2004 19626136 PMC2713747

      Not checked in the free run.

    • cites [45] Puelles L Pallial and subpallial derivatives in the embryonic chick and mouse telencephalon, traced by the expression of the genes Dlx-2, Emx-1, Nkx-2.1, Pax-6, and Tbr-1 J Comp Neurol 424 409 438 2000 10.1002/1096-9861(20000828)424:3<409::AID-CNE3>3.0.CO;2-7 10906711

      Not checked in the free run.

    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Study design and conductcitation

    The authors used only animals that did not sing after an overnight period of silence alone, because neural activity associated with singing causes changes in expression of thousands of genes in the song nuclei.

    “We used only animals that did not sing after an overnight period of silence alone, because neural activity associated with singing causes changes in expression of thousands of genes in the song nuclei (37, 42).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [37] Whitney O Science 346 1256780 2014 10.1126/science.1256780 25504732 PMC4359888

      Not checked in the free run.

    • cites [42] Jarvis ED Nottebohm F Motor-driven gene expression Proc Natl Acad Sci USA 94 4097 4102 1997 10.1073/pnas.94.8.4097 9108111 PMC20574

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Convergent changes in amino acid sequence among vocal-learning birds and mammals have recently been reported.

    “Likewise, convergent changes in amino acid sequence among vocal-learning birds and mammals have recently been reported (32, 33).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [32] Wang R thesis Duke University 2011 http://dukespace.lib.duke.edu/dspace/bitstream/handle/10161/5630/Wang_duke_0066D_10911.pdf?sequence=1

      Not checked in the free run.

    • cites [33] Zhang G Comparative genomics reveals insights into avian genome evolution and adaptation Science 346 1311 1320 2014 10.1126/science.1251385 25504712 PMC4390078

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Song-learning birds and humans are separated from a common ancestor by more than 68 million years among birds or 310 million years between birds and humans.

    “We examined whether complex behavioral and anatomical convergence is associated with molecular convergence in song-learning birds and humans, species separated from a common ancestor by more than 68 million years (among birds) or 310 million years (between birds and humans) (34–36).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [34] Jarvis ED Avian brains and a new understanding of vertebrate brain evolution Nat Rev Neurosci 6 151 159 2005 10.1038/nrn1606 15685220 PMC2507884

      Not checked in the free run.

    • cites [35] Kumar S Hedges SB A molecular timescale for vertebrate evolution Nature 392 917 920 1998 10.1038/31927 9582070 source

      Not checked in the free run.

    • cites [36] Jarvis ED Whole-genome analyses resolve early branches in the tree of life of modern birds Science 346 1320 1331 2014 10.1126/science.1253451 25504713 PMC4405904 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Songbird RA and HVC are part of a vocal motor pathway that controls production of learned vocalizations, whereas Area X and LMAN are part of a pathway that controls imitation and exploration of vocalizations.

    “The songbird RA and HVC are part of a vocal motor pathway that controls the production of learned vocalizations, whereas Area X and LMAN are part of a pathway that controls imitation and exploration of vocalizations (Fig. 1A) (40, 41).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites [40] Brainard MS Doupe AJ Translating birdsong: Songbirds as a model for basic and applied medical research Annu Rev Neurosci 36 489 517 2013 10.1146/annurev-neuro-060909-152826 23750515 PMC4130661

      Not checked in the free run.

    • cites [41] Jarvis ED Learned birdsong and the neurobiology of human language Ann NY Acad Sci 1016 749 777 2004 10.1196/annals.1298.038 15313804 PMC2485240

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    Avian RA and arcopallium are similar to mammalian layer 5 cells, but each vocal-learning species' RA analog has greater similarity to human laryngeal cortex regions relative to nonhuman primate motor cortex.

    “These findings demonstrate the similarity of avian RA and arcopallium to mammalian layer 5 cells, but a greater similarity of each vocal-learning species’ RA analog to human laryngeal cortex regions (which also contain layer 5 cells) relative to nonhuman primate motor cortex.”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationcitation

    The similarity to somatosensory cortex is not too surprising, considering the underappreciated overlap of cells with motor and somatosensory functions in both walls of the central sulcus.

    “The similarity to the somatosensory cortex is not too surprising, considering the underappreciated overlap of cells with motor and somatosensory functions in both walls (PrG_cs and PoG_cs) of the central sulcus (55).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [55] Nii Y Uematsu S Lesser RP Gordon B Does the central sulcus divide motor and sensory functions? Cortical mapping of human hand areas as revealed by electrical stimulation through subdural grid electrodes Neurology 46 360 367 1996 10.1212/WNL.46.2.360 8614495

      Not checked in the free run.

    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • May combine two claims.

    Interpretationown data

    The similarity between RA and the broader primary motor cortex in humans is due in part to similarity with its layer 5 neurons, and is separate from the similarities to LMC/dLSC within primary motor cortex.

    “These findings indicate that the similarity between RA and the broader primary motor cortex in humans is due in part to similarity with its layer 5 neurons, and is separate from the similarities to LMC/dLSC within primary motor cortex.”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.
  • Probably not a substantial claim: may not pass S2 (checkable).

    Interpretationreasoning

    The genes and their expression differences can serve as candidates that can lead us to the cause.

    “the genes and their expression differences can serve as candidates that can lead us to the cause”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Convergent gene expression for RA/LMC and Area X/putamen involves species separated from a common ancestor by 68 million to 310 million years, different vertebrate lineages, and many genes.

    “We find the convergent gene expression for RA/LMC and Area X/putamen remarkable in that they involve species separated from a common ancestor by 68 million to 310 million years (34–36), different vertebrate lineages, and many genes.”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [34] Jarvis ED Avian brains and a new understanding of vertebrate brain evolution Nat Rev Neurosci 6 151 159 2005 10.1038/nrn1606 15685220 PMC2507884

      Not checked in the free run.

    • cites [35] Kumar S Hedges SB A molecular timescale for vertebrate evolution Nature 392 917 920 1998 10.1038/31927 9582070 source

      Not checked in the free run.

    • cites [36] Jarvis ED Whole-genome analyses resolve early branches in the tree of life of modern birds Science 346 1320 1331 2014 10.1126/science.1253451 25504713 PMC4405904 source

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Our analyses do not show that the specializations discovered are functionally related to the projections of these neurons or to vocal learning.

    “our analyses do not show that the specializations we discovered are functionally related to the projections of these neurons or to vocal learning”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationown data

    The correlations are highly significant and predominantly driven by specialized expression of tens to hundreds of genes out of thousands, and for vocal-learning regions are supported by lack of such findings in vocal-nonlearning birds and primates.

    “Although the correlations we find at first glance may not seem strong, they are highly significant, are predominantly driven by the specialized expression of tens to hundreds of genes (depending on the level in the anatomical tree) out of thousands of genes, and for vocal-learning regions are supported by lack of such findings in vocal-nonlearning birds and primates.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationreasoning

    The match of Area X to human striatum and RA analogs to cortex enables specific hypotheses about which brain regions are analogous in song-learning birds and humans, and which regions in humans may be specialized for speech.

    “The match of Area X to the human striatum and of RA analogs to the cortex enables specific hypotheses about which brain regions are analogous in song-learning birds and humans, as well as which regions in humans may be specialized for speech.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    ROBO1 mutations cause dyslexia and speech disorders in humans.

    “ROBO1 mutations cause dyslexia and speech disorders in humans (76)”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [76] Fisher SE Francks C Genes, cognition and dyslexia: Learning to read the genome Trends Cognit Sci 10 250 257 2006 10.1016/j.tics.2006.04.003 16675285

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    The role of human striatum in speech acquisition and production has begun to be appreciated, but specific loci have not been as well defined as for cortex.

    “The role of the human striatum in speech acquisition and production has begun to be appreciated (57, 83), but specific loci have not been as well defined as for the cortex.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites [57] Brown S The somatotopy of speech: Phonation and articulation in the human motor cortex Brain Cognit 70 31 41 2009 10.1016/j.bandc.2008.12.006 19162389 PMC2873785 source

      Not checked in the free run.

    • cites [83] Simonyan K Horwitz B Jarvis ED Dopamine regulation of human speech and bird song: A critical review Brain Lang 122 142 150 2012 10.1016/j.bandl.2011.12.009 22284300 PMC3362661

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Parvalbumin is present at unusually high levels in RA neurons that project to vocal motor neurons in songbirds and in analogous layer 5 neurons of oral motor cortex of humans and chimpanzees, but not in projection neurons of adjacent arcopallium or in oral motor cortex of other primates or mammals.

    “the calcium-buffering protein parvalbumin has been noted (16) to be present at unusually high levels in the RA neurons that project to vocal motor neurons in songbirds and in the analogous layer 5 neurons of oral motor cortex (containing vLMC) of humans and chimpanzees, but not in the projection neurons of the adjacent arcopallium or in the oral motor cortex of other primates or mammals (79, 80).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites [16] Hara E Rivas MV Ward JM Okanoya K Jarvis ED Convergent differential regulation of parvalbumin in the brains of vocal learners PLOS ONE 7 e29457 2012 10.1371/journal.pone.0029457 22238614 PMC3253077

      Not checked in the free run.

    • cites [79] Wild JM Williams MN Suthers RA Parvalbumin-positive projection neurons characterise the vocal premotor pathway in male, but not female, zebra finches Brain Res 917 235 252 2001 10.1016/S0006-8993(01)02938-9 11640910

      Not checked in the free run.

    • cites [80] Sherwood CC Holloway RL Erwin JM Hof PR Cortical orofacial motor representation in Old World monkeys, great apes, and humans. II. Stereologic analysis of chemoarchitecture Brain Behav Evol 63 82 106 2004 10.1159/000075673 14685003

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    dLMC was only recently described.

    “the dLMC was also only recently described (63)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [63] Brown S Ngan E Liotti M A larynx area in the human motor cortex Cereb Cortex 18 837 845 2008 10.1093/cercor/bhm131 17652461 source

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Premisecitation

    A recent study argued avian pallium is equally and only weakly similar to mouse cortex, claustrum, or amygdala.

    “a recent study that used brain transcriptome profiles to argue that the avian pallium is equally and only weakly similar to the mouse cortex, claustrum, or amygdala (68)”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [68] Belgard TG Adult pallium transcriptomes surprise in not reflecting predicted homologies across diverse chicken and mouse pallial sectors Proc Natl Acad Sci USA 110 13150 13155 2013 10.1073/pnas.1307444110 23878249 PMC3740902

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    This lack does not mean that shared specializations of more specific regions within cortex do not exist.

    “does not mean that shared specializations of more specific regions within the cortex do not exist”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data

    Our ability to identify a stronger relationship to cortex was because we organized expression profiles as specialization trees, developed a dynamic programming algorithm requiring hierarchical anatomical alignment of brain similarities, and performed avian brain dissections using more precise laser capture microdissections.

    “We believe our ability to identify a stronger relationship to the cortex was because we organized the expression profiles as specialization trees, developed a dynamic programming algorithm requiring hierarchical anatomical alignment of brain similarities, and performed avian brain dissections using more precise laser capture microdissections.”
    Quote in the paperSubstantial claim
52 of 52
  • The authors state this as an assumption.

    The human premortem vocalizing state is unknown, but it is assumed not to prevent identification of convergent gene expression specializations as long as one species was silent, the genes are not sensitive to immediate vocalizing behavior, and human samples were collected 10 to 30 hours postmortem when many vocalizing-regulated gene expression changes have subsided.

    “We do not know the premortem vocalizing state of the humans, but did not expect this to prevent us from identifying convergent gene expression specializations as long as one species was silent (the birds), the genes are not sensitive to immediate vocalizing behavior, and the human samples were collected 10 to 30 hours postmortem, when many of the vocalizing-regulated gene expression changes have”
  • The paper relies on this without support.

    No examples of widespread convergence in gene expression patterns in brain regions involved in convergently evolved behavioral traits were known before this study.

    “However, we are not aware of examples of widespread convergence in gene expression patterns in brain regions involved in convergently evolved behavioral traits.”
2 of 2

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?

  • Studies on avian vocal learners have shown a convergent differential regulation of axon guidance genes of the SLIT-ROBO families in the pallial motor nucleus of the learning species, allowing for the formation of connections virtually absent in the brains of vocal non-learners.

    “Studies conducted on avian vocal learners89,90 have shown a convergent differential regulation of axon guidance genes of the SLIT-ROBO families in the pallial motor nucleus of the learning species, allowing for the formation of connections virtually absent in the brains of vocal non-learners.”

Cites

Flagged first, then by citations

1 of the 117 references couldn’t be matched to a known paper.

Research works printed but not linked: 1

  1. R. Wang thesis Duke University (2011); http://dukespace.lib.duke.edu/dspace/bitstream/handle/10161/5630/Wang_duke_0066D_10911.pdf?sequence=1.

Cited by

Sorted by citations

Authors
Pfenning, A. R., Hara, E., Whitney, O., Rivas, M. V., Wang, R., Roulhac, P. L., Howard, J. T., Wirthlin, M., Lovell, P. V., Ganapathy, G., Mountcastle, J., Moseley, M. A., Thompson, J. W., Soderblom, E. J., Iriki, A., Kato, M., Gilbert, M. T. P., Zhang, G., Bakken, T., Bongaarts, A., Bernard, A., Lein, E., Mello, C. V., Hartemink, A. J., & Jarvis, E. D.
Affiliations
  • Howard Hughes Medical Institute (US)
  • Oregon Health & Science University (US)
  • Duke University (US)
  • RIKEN Center for Brain Science (JP)
  • University of Copenhagen (DK)
  • Allen Institute for Brain Science (US)
  • Allen Institute for Brain Science, Seattle, WA 98103, USA.
Corresponding author
  • Pfenning, Andreas R.
  • Hartemink, Alexander J.
  • Jarvis, Erich D.
Venue
Science
Published
2014-12-12
Volume
346
Issue
6215
Pages
1256846-1256846
ISSN
  • 0036-8075 (ISSN-L)
  • 1095-9203
OpenAlex
W2052801529
Cited by
463
References
117
Citations by year
2026: 25 · 2025: 40 · 2024: 24 · 2023: 32 · 2022: 36 · 2021: 47 · 2020: 39 · 2019: 38 · 2018: 43 · 2017: 57 · 2016: 33 · 2015: 44 · 2014: 4 · 2013: 1
FWCI
42.23
Type
journal article
Language
en
Open access
Locations
  • Science (publishedVersion)
  • PubMed (publishedVersion)
  • Europe PMC (PubMed Central) (acceptedVersion)
  • PubMed Central (acceptedVersion)
  • DukeSpace (Duke University) (submittedVersion, cc-by-nc-nd)
  • eSpace (Curtin University) (submittedVersion)
  • Research at the University of Copenhagen (University of Copenhagen) (submittedVersion)
Funders
  • Howard Hughes Medical Institute
  • National Institutes of Health (DP1OD000448)
  • National Institute on Deafness and Other Communication Disorders (R21DC007478, R01DC007218)
  • National Institute of General Medical Sciences (R24GM092842)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Neuroscience and Music Perception
  • Social Sciences › Cultural Studies › Language and cultural evolution
Keywords
vocal learning, human speech, convergent evolution, brain transcriptome, gene expression, song nuclei, striatum, motor control, brain connectivity, avian vocal learning
MeSH terms
Adult, Animals, Birds, Brain, Brain Mapping, Corpus Striatum, Gene Expression Regulation, Humans, Learning, Male, Motor Cortex, Neural Pathways, Species Specificity, Speech, Transcription, Genetic, Vocalization, Animal, Evolution, Molecular, Finches, Transcriptome

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.

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