Revisiting the Hypothesis of Language Retrogenesis From an Evolutionary Perspective

Abstract

Objective:
In this article, we reexamine the hypothesis of language retrogenesis, that is, the assumption that language change over healthy ageing mirrors, albeit inversely, language acquisition by the child. We additionally question whether this inverse pattern can as well be observed at the cognitive and neurobiological levels, and whether it can be informative (and a consequence, in fact) of how language evolved in humans.

Method:
We compare the language strengths and weaknesses signifying language acquisition and its eventual decay in healthy ageing. We further compare age-related cognitive and neurobiological readjustments during each of these two developmental stages, with a focus on brain areas involved in language processing. Finally, we delve into the evolutionary changes experienced by these areas.

Results:
We present evidence supporting the hypothesis of retrogenesis in two domains of language: the lexicon (lexical access, understanding of nonliteral meanings, and resolution of lexical competition) and syntax (understanding and production of complex sentences). Additionally, we show evidence that the brain areas supporting these complex tasks are late-myelinated in childhood and early-demyelinated during ageing. Finally, we show that some of these areas (such as the inferior frontal gyrus) are phylogenetically newer.

Conclusions:
Language acquisition in children and language degradation/loss in healthy ageing follow the principle of retrogenesis, but mostly in domains that are cognitively demanding and that depend on recently evolved brain devices. Putting this differently, the components of language that emerged more recently appear to be more, and earlier, affected during ageing, as well as developed later over childhood. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Full text

Neuropsychology
Revisiting the Hypothesis of Language Retrogenesis From an Evolutionary
Perspective
Antonio Benítez-Burraco and Olga Ivanova
Online First Publication, February 2, 2023. https://dx.doi.org/10.1037/neu0000886
CITATION
Benítez-Burraco, A., & Ivanova, O. (2023, February 2). Revisiting the Hypothesis of Language Retrogenesis From an
Evolutionary Perspective. Neuropsychology. Advance online publication. https://dx.doi.org/10.1037/neu0000886

Revisiting the Hypothesis of Language Retrogenesis From an
Evolutionary Perspective
Antonio Benítez-Burraco
1
and Olga Ivanova
2
1
Department of Spanish, Linguistics and Theory of Literature (Linguistics), Faculty of Philology, University of Seville
2
Spanish Language Department, Faculty of Philology, University of Salamanca
Objective: In this article, we reexamine the hypothesis of language retrogenesis, that is, the assumption that
language change over healthy ageing mirrors, albeit inversely, language acquisition by the child. We
additionally question whether this inverse pattern can as well be observed at the cognitive and neurobio-
logical levels, and whether it can be informative (and a consequence, in fact) of how language evolved in
humans. Method: We compare the language strengths and weaknesses signifying language acquisition and
its eventual decay in healthy ageing. We further compare age-related cognitive and neurobiological
readjustments during each of these two developmental stages, with a focus on brain areas involved in
language processing. Finally, we delve into the evolutionary changes experienced by these areas. Results:
We present evidence supporting the hypothesis of retrogenesis in two domains of language: the lexicon
(lexical access, understanding of nonliteral meanings, and resolution of lexical competition) and syntax
(understanding and production of complex sentences). Additionally, we show evidence that the brain areas
supporting these complex tasks are late-myelinated in childhood and early-demyelinated during ageing.
Finally, we show that some of these areas (such as the inferior frontal gyrus) are phylogenetically newer.
Conclusions: Language acquisition in children and language degradation/loss in healthy ageing follow the
principle of retrogenesis, but mostly in domains that are cognitively demanding and that depend on recently
evolved brain devices. Putting this differently, the components of language that emerged more recently
appear to be more, and earlier, affected during ageing, as well as developed later over childhood.
Key Points
Question: We revisit the hypothesis of language retrogenesis from a linguistic, cognitive, and
neurobiological perspectives. Findings: We conclude that a retrogenetic pattern can be observed in
healthy ageing, in particular, with regards to the most cognitively demanding aspects of language, like
complex syntax or complex semantics. Importance: Language changes in healthy ageing may be
explored as a proxy to language evolution. Next Steps: Genetic, cross-cultural, and cross-linguistic
studies are essential to confirm these parallels between language ontogeny and phylogeny in humans.
Keywords: language ageing, language acquisition, language complexity, language retrogenesis
Originally based on the “law of regression”proposed by Théodule
Ribot (1887),thehypothesis of retrogenesis argues that degen-
erative processes mirror, albeit inversely, the order of acquisition
of functions or abilities in child normotypical development
(Reisberg et al., 1999). The hypothesis of retrogenesis is thus
mainly used to explain cognitive and neurofunctional changes
in pathological ageing. Neurobiological retrogenesis broadly as-
sumes that developmentally late-myelinating fibers in the brain are
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Antonio Benítez-Burraco https://orcid.org/0000-0003-4574-5666
Olga Ivanova https://orcid.org/0000-0002-9657-5380
This research was supported by Grant PID2020-114516GB-I00 funded by
Spanish Ministry of Science and Innovation/Spanish State Investigation
Agency/10.13039/501100011033 (awarded to Antonio Benítez-Burraco).
The authors have no conflicts of interest to declare.
Antonio Benítez-Burraco and Olga Ivanova conceived the article, per-
formed the data analyses, and wrote and approved the final article.
Geolocation: Seville, Spain, Europe.
Antonio Benítez-Burraco played lead role in funding acquisition and equal
role in conceptualization, investigation, methodology, writing of original
draft and writing of review and editing. Olga Ivanova played equal role in
conceptualization, investigation, methodology, writing of original draft and
writing of review and editing.
The research conducted for the article relied on previously published
data by others and available data sets, hence no ethics approval was
required.
The material used in the analyses is presented as a list of bibliographical
references at the end of the article. A previous version of this article (now
significantly improved) has been posted as a preprint to PsyArXiv: https://
www.psyarxiv.com/6khb5/.
Correspondence concerning this article should be addressed to Antonio
Benítez-Burraco, Department of Spanish, Linguistics and Theory of
Literature (Linguistics), Faculty of Philology, University of Seville,
C/Palos de la Frontera s/n. 41004 Sevilla, Spain. Email: abenitez8@us.es
Neuropsychology
© 2023 American Psychological Association
ISSN: 0894-4105 https://doi.org/10.1037/neu0000886
1

more susceptible to early degeneration (i.e., demyelination;
Brickman et al., 2012), resulting in reduced intrinsic connectivity
and processing efficiency in age-related and pathological condi-
tions (Kavroulakis et al., 2021). Cognitive retrogenesis presup-
poses that cognitive functions are more susceptible of early decline
in ageing if they expand later in childhood (cf. Simoes Loureiro
& Lefebvre, 2016). Drawing from the dependency of cognitive
regression on neurobiological retrogenesis, one can expect lan-
guage retrogenesis to follow the hierarchical pattern too, with
ageing or, more commonly, demented speakers generating more
immature and early acquired language patterns (see Emery, 2000;
Gayraud et al., 2019;Haak, 2002).
While the hypothesis of retrogenesis is not unanimously accepted
for language (see Lust et al., 2015;Moos, 2011, for opposing view-
points), many studies argue for a hierarchical language decline in
pathological ageing (Gayraud et al., 2019;Kim et al., 2011;Rogers &
Lasprilla, 2006;Simoes Loureiro & Lefebvre, 2016). Indeed, the
hypothesis of language retrogenesis is mainly applied to and tested on
dementias, above all Alzheimer’s disease (AD; Ahmed et al., 2017;
Kim et al., 2011;Mello et al., 2008;Moos, 2011;Reisberg et al.,
2002), but also semantic dementia (SD; Pozueta et al., 2020). Despite
limited data available, retrogenesis is suggested to affect the lexical
level (impacting mostly on nouns), with later-acquired vocabulary
being lost first (Kim et al., 2011), and later-acquired, subordinate-level
semantic knowledge being more easily disrupted in demented speak-
ers (Pozueta et al., 2020;Simoes Loureiro & Lefebvre, 2016). In
addition to the lexical–semantic level, syntactic simplification is also
suggested to follow a retrogenetic pattern in dementia (Haak, 2002),
though this assumption is not undisputedly accepted either (see Lust
et al., 2015, for discussion).
Two rationales can explain the so-called imperfection of the
retrogenetic model of demented language (Caramazza, 1994, for
the pioneering work challenging the hypothesis of retrogenesis).
First, as we will discuss below, different levels/domains of language
can be differently affected by retrogenesis. Accordingly, language
levels with a more procedural embedding can be more immune to
retrogenesis, like phonology, while others might follow the retro-
genetic model more closely, like syntax. Second, deviations from
the retrogenetic model can result from intrinsic variation across
age groups typically considered for testing this model (children vs.
elderly). A good example is semantic categorization, which is
steadily conditioned by the cultural context of language acquisition:
only when its effects are disregarded (e.g., when taxonomic asso-
ciations are excluded), the pattern of semantic deterioration repli-
cates the retrogenetic principle (see Mello et al., 2008, for insightful
data in this respect).
To what extent language change in neurotypical ageing echoes the
retrogenetic pattern of language in dementia is still a topic to explore.
It is true that healthy ageing itself is rarely associated with a straight
language impairment: recent work (e.g., Peelle, 2019) points rather to
a cognitive (and, thus, language-specific) readjustment for successful
language processing and production in ageing. However, there is also
evidence that the loci of the neuroanatomically heightened vulnera-
bility in AD correspond to late-developing brain areas in children and
early-degenerating brain areas in healthy ageing (Douaud et al.,
2014). Additionally, preliminary studies support the idea that AD-
induced cognitive (and language) retrogenesis impacts neocortical-
related functions (Rubial-Álvarez et al., 2013), suggesting that the
earliest AD-driven disruptions could affect the most recently
acquired cognitive functions (Arendt et al., 2017). Studies on
changes in white matter (Brickman et al., 2012), reduction in
myelin content (Papadaki et al., 2019) and functional intrinsic
connectivity (Kavroulakis et al., 2021) support the view that
healthy ageing could be a candidate to neurocognitive retrogenesis.
A reason is that, for example, studies on white matter integrity in
healthy and pathological ageing suggest that the same (retroge-
netic) neuroanatomical changes can be observed across all ageing
spectrum, with AD showing more pronounced alterations com-
pared to healthy elderly (Gao et al., 2011). Likewise, cognitive
decline in healthy ageing and dementias seems to share similar
pathophysiological mechanisms (Gonzales et al., 2022). It is, thus,
possible that (some of) the decline pathways in the two types of
ageing, particularly with regard to language, could also be similar,
supporting the hypothesis of language retrogenesis for neurotypi-
cal cases too.
Accordingly, our own wording of the ontogenetic language
retrogenesis hypothesis -the focus of the present article claims
that the language patterns acquired later in children are disrupted
earlier in healthy ageing, with the addenda that these patterns could
be regarded as phylogenetically more recent. As we will argue, these
evolutionary considerations are crucial for giving support to the
retrogenetic hypothesis. The reason is that recently evolved brain
and cognitive components are more sensitive to ontogenetic damage
because of the lack of compensatory mechanisms, which would
usually develop after a prolonged period of selection and evolution
(see Benítez-Burraco & Boeckx, 2014;Benítez-Burraco & Murphy,
2019,orBenítez-Burraco, 2020, for discussion). If the hypothesis of
language retrogenesis is confirmed, it may have a major impact on
our understanding of the nature and evolution of human language.
Specifically, it would give additional support to the view that
different language components are subject to different degrees of
cognitive complexity and, ultimately, brain-processing demands,
with the more complex aspects evolving later and with the less
complex showing more evolutionary continuity with other species.
Since language does not fossilize, we need to rely on indirect
evidence to infer the historical image of human languages and
the evolution of our ability to learn and use them.
The objective of this article is to check the plausibility of the
ontogenetic language retrogenesis hypothesis and to explore its
potential role as a lens to the evolution of language in humans.
To this end, we aim to provide an exhaustive comparative view
between language development in the child and language changes in
healthy ageing. We will also consider cognitive and neurobiological
changes during development and ageing, assuming that all these
changes can be used as a basis for reconstructing language evolution.
Here, a few points are worth clarifying. First, we are not claiming
that ontogeny exactly recapitulates phylogeny. That said, when
changes in language structure and its use during growth are consid-
ered together with the developmental changes in brain anatomy
and cognitive functionality, robust evolutionary findings can be
achieved. As noted by Pattabiraman et al. (2020), our species-
specific cognitive abilities arose in part from changes in the organi-
zation and function of preexisting neural devices, and these changes
account for distinctive aspects of our neurodevelopment, but also for
our predisposition to suffer from ontogenetic damage and neurode-
generative diseases.
Second, we base our approach on viewing the language as a
specialized cognitive ability resulting from the interaction between
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2BENÍTEZ-BURRACO AND IVANOVA

the brain areas otherwise performing basic types of computations
that are recruited for different cognitive abilities, in the spirit of
Poeppel and Embick (2005), among many others. This ability is put
into use for fulfilling many different functions, from thinking to
communicating to socializing. We further consider languages as
specific codes in which this general ability is displayed in different
human groups, with their distinctive properties resulting in part from
the properties of language as a cognitive and behavioral component
of the human phenotype, but also from environmental and cultural
constraints. Finally, we acknowledge that language evolution,
including the evolution of modern languages, is the combined
outcome of both a biological process, impacting our brain and
behavior, and a cultural process, responsible for distinctive language
properties resulting from its transmission and use. Specifically, we
support the view that biological changes that brought about our
species also favored the creation of the cultural niche that enabled
the sophistication of languages through a cultural mechanism (see
Benítez-Burraco & Progovac, 2020, for a detailed discussion). We
find this view in line with eco–evo–devo theories in biology,
according to which organisms evolve as a result of the interactions
between their genes, their developmental paths, and the environ-
ments in which they live (Gilbert et al., 2015;Segovia-Martín&
Balari, 2020).
We structure our article as following. First, we brieflyarguewhy
early stages of language development in children, as well as initial
stages of language deterioration during ageing, can be regarded as
“degraded”forms of language, following Bickerton (1990) and
Jackendoff (1999), among others, and eventually, can be viewed
as a proxy to phylogenetic stages of language evolution. Second,
given that stages and milestones of language development in
children have been explored more profoundly, and generally
show greater in-group consistency, we provide an in-depth review
of the current developments in language changes in healthy
ageing. Third, we provide a detailed comparison of language
changes in healthy ageing and the milestones of language acqui-
sition in children. Next, we provide a brief comparison between
the anatomical and functional changes in the brain during language
acquisition in children and the corresponding changes during
healthy ageing, searching for parallels between language devel-
opment and its later readjustment at the neurobiological level.
While assessing similarities and differences between the two
processes, we do not recur to a systematic revision of literature,
but, rather, rely on qualitative, scoping-close approach, synthesiz-
ing existing evidence as a form of addressing far-reaching ques-
tions for novel insights (Cook, 2019;Munn et al., 2018). To avoid
evidence suppression bias, we also discuss arguments against the
theory of retrogenesis and our understanding of it. Finally, we
discuss our results in light of the current theories on evolutionary
changes in neuronal substrate of language, and, more generally,
current hypotheses on language evolution in the species. This will
make us reconsider, once again, the idea that language ontogeny
recapitulates language phylogeny. We will do so from the evo–
devo perspective, according to which evolution is fueled by
changes in developmental processes; that is, that ontogeny creates
phylogeny. We will conclude that our approach enables us to use
the model of language acquisition by a child versus language
changes in healthy ageing as informative proxies to specificstages
of language evolution.
Language Retrogenesis and “Degraded”Form of
Language
Multiple efforts have been made to shed light on the evolution of
the human language faculty (Fitch, 2017), with one of the most
interesting, yet less explored sources of evidence addressing what
Bickerton (1990) and Jackendoff (1999) labeled as the “degraded”
forms of language. This term is used to refer to different types of
structurally simplified and functionally reduced linguistic systems,
which may be found in speakers with language deficits resulting
from developmental or acquired disorders, or speakers with incom-
plete proficiency in the language that is being acquired as L1 or
learned as L2. The term also encompasses language systems with
limited expressive possibilities, such as lingua franca or pidgins.
Despite unclarity on the scopes and limits of the so-called
“degraded”forms of language, with the complexity of a linguistic
system being a matter of degree, one may assume that among such
we can safely include any linguistic system that lacks some/most of
the structural features found in natural languages and that does not
allow for complete functional communication. Accordingly, candi-
dates for the “degraded”forms of language (and, in turn, for proxies
of the earlier stages of the language evolution) can include any of the
following varieties, most of which were proposed by Jackendoff
(1999) with reference to Bickerton (2016) and Goldin-Meadow and
Mylander (1990): (a) pathological language instances, either result-
ing from ontogenetic damage or acquired impairment, for example,
agrammatic aphasias; (b) pidgin languages and other restricted
languages; (c) deaf children homesigns in nonsigning families;
and (d) (initial stages in the development of the) child language.
Still, there are disagreements about these “degraded”forms of
language. For example, Botha (2006) critically discussed the useful-
ness of these linguistic systems for language evolution studies,
whereas other authors raised concerns about certain specific types
of such candidates; see, for example, Slobin (2005), for child
language as a “degraded”form of language. However, over the
years, a few more candidates for “degraded”forms of language were
suggested. Two further cases are interlanguages developed by L2
learners (Roberge, 2009) and early stages in the language learning
process by computational models/artificial intelligence (Steels &
Szathmáry, 2018).
As noted, “degraded”forms of language share many distinctive
properties. For instance, the so-called Basic Variety, in which most
L2 learners fossilize (Klein & Perdue, 1997), features many of the
structural properties found in pidgins, including the absence of
inflectional morphology or subordination. At the same time, these
are features that are acquired later by children and are more easily
lost in patients with different aphasia syndromes, as suggested by
studies on milestones in child language development (Luinge et al.,
2006), language profiling in aphasia syndromes (Clark &
Cummings, 2003), or pattern models of language decline in demen-
tia (cf. Emery, 2000).
Therefore, some structural aspects of language (as well as some of
its functions) can be hypothesized to be more complex and more
demanding in terms of computational and operational rules and
meanings. As such, more complex elements of language structure
may end up taking more time, and/or bigger effort, and/or richer
learning/usage environment to be acquired. Ultimately, more com-
plex aspects of language may be processed less efficiently and/or
demand more cognitive/brain resources, and would, thus, be more
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LANGUAGE RETROGENESIS AND LANGUAGE EVOLUTION 3

vulnerable to any type of decline, both neurotypical and pathologi-
cal. As Jackendoff and Wittenberg (2014) assume with regard to
grammar, a language processor has multiple pathways at its dis-
posal, and when difficulties arise and the most complex ones (i.e.,
full grammar) fail, the simpler ones (i.e., linear grammar) may still
be available. If this assumption is correct, the neurobiological and
cognitive readjustment and deterioration in healthy ageing would
make more complex language structures more difficult to process
and produce, and, thus, more eligible to be replaced by simpler ones.
Indeed, this pattern is observed in several age-driven neurodegen-
erative conditions, including AD (Kim et al., 2011;Simoes Loureiro
& Lefebvre, 2016) and SD (Pozueta et al., 2020). In the same line,
some early works on language in ageing found that older adults tend
to avoid structures imposing higher memory load (e.g., subject–
relative sentences, complex sentences with multiple-embedded
clauses, etc.; Kynette & Kemper, 1986).
One intriguing possibility is that these more complex aspects of
language had also evolved more recently, and so the consideration
of language changes in ageing would significantly enrich the study
of language evolution. Within the domain of evolutionary linguis-
tics, the common features exhibited by the “degraded”forms of
language are equated, as noted, to the putative features exhibited
by the (proto)languages spoken by our species in the distant past
and/or by other hominin species in our lineage. Explicit claims
about such possibility mostly concern impaired language in adult
speakers. For instance, Code (2011) relied on his studies of
nonfluent aphasia resulting from damage in the left inferior frontal
brain areas to suggest that lexical speech automatisms in diseased
patients could mimic a prefull syntax stage of language evolution.
Interestingly, Code also argued that the recovery of aphasic speech
mimics [the early stages of] language evolution in our species.
Likewise, Ardila (2006,2015) took aphasia-induced language
changes as a basis for his view of language origins, suggesting
the two-stage evolution process of the human language: the
selectional, or lexical–semantic stage (as inspired by Wernicke’s
aphasia), and the sequential, or grammatical stage (as inspired by
Broca’s aphasia). Underlying this kind of hypotheses is a view that
language evolves from simpler to more complex forms, and that a
complex language demands more sophisticated cognitive and
behavioral abilities to be mastered; abilities that have only devel-
oped fully in our contemporaries. In their seminal article, Klein and
Perdue (1997) disagreed with the idea that their basic variety could
be assigned to a protolanguage distinct from the “fully-fledged”
languages, but acknowledged that it could be argued to exhibit
the core attributes of the human language capacity. Accordingly,
the missing features would be regarded as secondary and, thus,
as developed later in phylogeny. Likewise, Jackendoff and
Wittenberg (2014) noted that simpler forms of language that could
have been useful for hominins prior to the emergence of the full
language were not replaced but, rather, elaborated upon over the
course of language evolution.
Finally, despite being a catchy cover term for a set of interesting
language phenomena pointing to differences in complexity across
linguistic systems, the assumption that these “degraded”forms of
language are genuine proxies of the earlier stages of language
evolution needs to be reexamined under the light of the theory of
evolution. One crucial question is whether the “degraded”forms of
language can be regarded as real atavisms. True atavisms do exist in
biology (Hall, 2010, for discussion), and real reversions to ancestral
states can occur, but only if ancestral developmental programs are
preserved (but silenced) in the derived population (Cabej, 2012).
Interestingly, genes that were positively selected in our species are
enriched in candidate genes for conditions like schizophrenia or
autism spectrum disorder (ASD; see Srinivasan et al., 2016, and
Polimanti & Gelernter, 2017, respectively). These conditions entail
language deficits that resemble many of the features found in the
“degraded”forms of language, like the absence of inflectional
morphology or high-level syntax (see Tager-Flusberg & Joseph,
2003, for ASD). This suggests that the evolutionary changes that
resulted in our distinctive cognitive phenotype, including language,
brought about an increased susceptibility to cognitive disorders,
including language impairments. Conversely, damages to the lan-
guage phenotype can be observed during abnormal development
because of this common genetic signature. This is seemingly
explained by the reduced resilience of recently evolved neuronal
networks supporting them (see Toro et al., 2010, for ASD, and also
Pattabiraman et al., 2020). Still, some caution should be exercised.
One important aspect to be considered is that because this kind
of disorders are brought about by some type of damage—either
genomic (in the case of developmental language disorders) or
neurobiological (in the case of acquired language disorders)—it
might be difficult to differentiate between truly atavistic features and
maladaptive symptoms resulting from the damage. In developmen-
tal disorders, what we observe is a brain that is developing differ-
ently to accommodate certain genetic alteration. In acquired
disorders, it is a neurotypical brain that accommodates neurological
damage that occurred after its development has completed. In other
words, one must be cautious when assuming that these pathological
function losses truly mirror the putative functional gains in our
ancestors (see Benítez-Burraco & Boeckx, 2014, for further discus-
sion). For these reasons, we do not consider here brain disorders-
induced language changes, but instead focus on changes associated
with neurotypical growth and ageing.
Language and Cognitive Changes in Healthy Ageing
Significant efforts have been made to trace trajectories of cogni-
tive (Glisky, 2007;Salthouse, 2019), language (Burke & Shafto,
2004,2007;Peelle, 2019;Thornton & Light, 2006), and cognitive-
language changes (Shafto & Tyler, 2014) that tend to occur
throughout healthy ageing. In general, cognitive decline leads to
age-related language changes, mostly consisting in structural sim-
plification, functional reduction, and poorer processing abilities.
Also, there is a tendency for certain language functions to degrade
first, for others to follow-up later, and for some others to be
considerably preserved (see Burke et al., 2000, for an insightful
discussion). Although the relationships between ageing brain, age-
ing cognition, and ageing language are dynamic and can be affected
by multiple internal and external factors, we can use the studies at
hand to draw a clearer picture of age-related patterns of neurocog-
nitive and language changes. This would be incomplete without
mentioning the following four key findings.
First, ageing has an impact on broad spectrum of cognitive abilities
that are crucially involved in language processing, such as executive
function, attention, learning ability, concentration, working memory,
and inhibitory control (Abrams & Farrell, 2010). Yet, even in
the notable cases of prolonged language preservation in ageing
(see Cohen et al., 2019;Pistono et al., 2021), ageing language
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4BENÍTEZ-BURRACO AND IVANOVA

appears to involve a greater degree of variability in cognitive
functions comparing to younger speakers (Harada et al., 2013;
Leal & Yassa, 2019). Among other things, ageing brings about
changes in speech that primarily stem from ongoing changes in motor
execution (Tremblay et al., 2018) as well as from anatomical shifts
that impact voice quality, speed, and pitch (Tucker et al., 2021). As
argued above, in this article we focus on those cognitive changes with
a greater impact on language structure, particularly, if this impact is
also supported by evidence from neurodegenerative conditions.
By this reason, we do not discuss changes in the language usage
patterns (i.e., pragmatics) either, although there already exist
insightful discussions around ageing pragmatics (Bambinietal.,
2021). Pragmatics focuses on the interpretation of any human, not
only verbal behavior (Carston, 2002). Pragmatic competence is
indeed reported to be seriously compromised in acquired language
disorders, specifically in dementia (Cummings, 2007), but, as
noted, it depends on calculations other than purely linguistic,
mainly the theory of mind and mental time travel, so that it is
closer to a “social competence”(Jagoe, 2017). Additionally, prag-
matic involution does not seem entirely systematic in the specific
case of pathological ageing, particularly, because speakers with
dementia recur to interactional compensatory strategies that can
enable successful communication despite important underlying
cognitive and language deficits (Guendouzi, 2013;Huang et al.,
2022). Also, patterns of socialization, cultural factors, and behav-
ioral schemes have a direct implication in such involutionary
processes in pragmatics.
Second, a very rough picture of the changes in language structure
in healthy ageing suggests that there exists a dissociation between
production, which is more disrupted, and comprehension, which is
(apparently) more preserved (Abrams & Farrell, 2010;Burke &
Mackay, 1997).
Third, the same rough picture also suggests that more complex
language activities seem to be disrupted earlier during ageing. Here,
by more complex we refer to more intricate combinations, more
structured sequences, deeper hierarchical constructs, more sophisti-
cated or elaborated propositional contents, and/or advanced concep-
tuality. More complex language tasks demand more advanced and
more precise cognitive abilities, such as enhanced memory or infer-
ential competence, high-level processing capacity, or well-developed
perceptual-motor abilities and reasoning (Christiansen & Chater,
2008, for an overview of cognitive and biological prerequisites of
language complexity).
Finally, during healthy ageing, lexical and syntactic language
tasks that can be regarded as complex according to the previous
characterization (e.g., nonliteral meanings, semantically saturated
units, embedded subordinate clauses, and the like) appear to expe-
rience greater disruption compared to more accessible and less
cognitively demanding aspects of language. These specific levels
are the focus of our analysis in this article.
By contrast, we exclude from our analysis other language do-
mains, particularly, phonology and morphology. One reason is that
the latter are less tractable in terms of simplicity/complexity (spe-
cifically, in comparison with other “degraded”forms of language).
More importantly, in dementia, phonology and morphology seem to
remain more robust. Indeed, pure phonological errors in patients are
scarce and unsystematic, with phonological mismatches mainly
resulting from disrupted semantic representations, which display
as phonemic blending or transpositioning (see Hoffman et al., 2009
or Meteyard & Patterson, 2009 forSD),orfromdeficit in inhibiting
other phonological representations, which displays as phonologi-
cal interferences (see Faust et al., 2004 forAD).Overall,most
research point to a secondary, sporadic, or atypical alteration of
phonology in the language of people with dementia (see Benedet
et al., 2006 for SD; Croot et al., 2000, for atypical AD). In fact,
there is some evidence that speakers can rely more on phonology
as lexical–semantic knowledge degrades (see Reilly et al., 2007
or Reilly & Peelle, 2008, for SD). Morphology shows a similar
pattern, with performance in tasks evaluating this domain being at
the ceiling if there are no semantic constraints; this is true even for
languages with a rich morphology, as Hebrew (see Kavéet al.,
2012, for SD and Kavé& Levy, 2003, for AD). Difficulties are
mainly observed for irregular word forms (see Auclair-Ouellet,
2015 or Walenski et al., 2009, for AD). Morphological shifting
and mismatching seem to be due to semantic interference and/or
reduced access to semantic networks (see Nikolaev et al., 2019 and
Simonsen et al., 2004, for AD). Finally, there is also an evolution-
ary reason for excluding phonology and morphology: these
are the domains of language that exhibit more continuity with
animal communication, with many animal calls showing human-
like sound structures (Samuels, 2015) and with some of them
having signs of morphological compositionality (Arnold &
Zuberbühler, 2006), although it is also true that both phonology
and morphology have been subject as well to evolution in the
human clade (e.g., Fitch, 2019 on phonology).
Language Production
As mentioned earlier, language production is the most affected
language domain during ageing, with lexical (word selection and
production) and syntactic (formation of complex sentences) levels
taking the biggest hit.
Lexical productive difficulties in ageing are mainly centered
around the global phenomenon of word-finding. Healthy elderlies
are reported to struggle when accessing mental lexicon as measured
through naming (Melvold et al., 1994), to exhibit word-finding
difficulties resulting in frequent circumlocutions (Glisky, 2007),
and to expose tip-of-the-tongue problems during word retrieval
(Burke & Mackay, 1997). These difficulties are particularly notice-
able when accessing and producing proper names (Ouyang et al.,
2020), mainly due to their specific semantic structure when compared
to the common words (Juncos-Rabadánetal.,2010). Among com-
mon names, problems are initially observed with infrequent and long-
inactive words due to insufficient priming and poorer semantic
connections (Barresi et al., 2000). This suggests that more complex
and less semantically connected words are among the first candidates
to be lost. Overall, incidental word-finding problems in ageing point
to a general difficulty for accessing and recovering word-forms
(Burke et al., 2000). Experimental evidence is indicative of a
disruption of the elderly’s capacity to adjust lexical concepts to their
respective phonological forms (Barresi et al., 2000;Burke & Mackay,
1997;Tremblay et al., 2018). This would explain language produc-
tion deficits, mainly dysfluencies, word repetitions, gap filling, and
phonemic omissions (Burke & Graham, 2012), but also the observed
positive effect of phonological priming (Ouyang et al., 2020)and
context facilitation effects (Burke et al., 2000)onwordretrieval
in ageing.
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LANGUAGE RETROGENESIS AND LANGUAGE EVOLUTION 5

These word-finding problems seem to boil down to neurobiolog-
ical changes in the elderly brain. Although we address these changes
in more detail in the next section, it is worth mentioning that the tip-
of-the-tongue effects are hypothesized to result from reduced gray
matter in the left insula, known to be implicated in phonological
production (Shafto et al., 2007). A reduction of the left prefrontal
cortex and the temporo-occipital regions, resulting in increases in
the insular regions activity (Manenti et al., 2013), and/or a reduction
in dopaminergic and cholinergic activity in selected areas of the
brain (Albert, 1990), are also related to this phenomenon.
At the same time, older adults feature an extremely extensive
vocabulary, as vocabulary size increases over the life span (Glisky,
2007;Gollan & Goldrick, 2019). They also exhibit advanced lexical
sophistication and semantic expertise (Spreng & Turner, 2019),
deeply relying on crystallized knowledge (Rabaglia & Salthouse,
2011), which is argued to compensate for reduced cognitive control
observed in ageing (Amer et al., 2016).
In the domain of syntax, older adults generally tend to produce
shorter and less complex sentences. Ageing results in the preference
for coordinated and right-branching syntactic constructions
(Kemper et al., 2001), and in avoidance of left-branching structures
and sentences with dense propositional contents (Rabaglia &
Salthouse, 2011). Simpler sentences in ageing are characterized
by vaguer terms and longer gaps (Shafto & Tyler, 2014). These
changes are hypothesized to result from reduction in available
working memory (Kempler et al., 1998), and, generally, to be
privileged by lower memory demands (Venneri et al., 2018),
seemingly boiling down to decreased functional connectivity in
syntax-supporting neural networks, mainly in left pars opercularis
(Antonenko et al., 2013). That said, elderly speakers seem to
maintain their ability to follow syntactic priming and lexical boost,
and, ultimately, to activate syntactic representations fully (Hardy
et al., 2017). Because of that, several authors have suggested that
syntactic anomalies in healthy ageing result mostly from lexical
difficulties of sorts highlighted above, rather than from a specific
impairment of syntactic processes (Hardy et al., 2020).
Language Comprehension
As mentioned above, when compared to language production,
language comprehension abilities appear to be better preserved in
the elderly, likely because of their richer linguistic knowledge and
command for procedural rules (hence the notable preservation of
morphology) as opposed to other cognitive declines (Wingfield
et al., 2015). Yet, older adults also exhibit difficulties in different
domains of language comprehension.
Vocabulary comprehension in healthy ageing is usually charac-
terized as spared. However, two specificdeficits mimicking lexical
production are observed in comprehension too: these are lexical
competition and difficulties with integrating contextual information.
The former implies ageing speakers to be more prone to activate
neighboring nontarget words, with negative impact on accessing
infrequent words and words with multiple phonological neighbors
(DeDe & Knilans Flax, 2016). Consequently, ageing people gradu-
ally lose their ability to predict word meaning from the contextual
cues (Wlotko et al., 2010). In general, context is a burden for older
adults: word comprehension problems are more pronounced in
noisy contexts (Tucker et al., 2021), and resolution of lexical
ambiguity becomes more costly as the context becomes more
semantically impoverished (Lee & Federmeier, 2011). Interestingly,
idiomatic expressions, such as proverbs, can be interpreted literally
because of higher executive demands (Uekermann et al., 2008). This
is in line with a decline in the ability to comprehend conventional
metaphors, even in situational contexts (Sundaray et al., 2018).
Overall, these deficits can be explained, at least partially, in terms of
age-related general inhibition deficit that results in lower inhibitory
control of lexical information (Mattys & Scharenborg, 2014).
Executive deficits can also account for these problems, since
demands are higher for achieving correct interpretation of utterances
(see Uekermann et al., 2008).
Syntactic comprehension problems in healthy ageing can mimic
vocabulary comprehension disruption. In general, older adults show
difficulties with understanding sentences with complex syntactic
structures or those that are syntactically ambiguous (Abrams &
Farrell, 2010). Demanding sentences, such as those containing long-
distance dependencies or embedded clauses, can lead to longer
online processing times for older speakers and, ultimately, to
comprehension problems (Caplan et al., 2011;Wingfield &
Grossman, 2006). Sentences with noncanonical word order can
also cause comprehension disruptions (DeDe & Knilans Flax,
2016), just as sentences with agreement errors (Poulisse et al., 2019).
Overall, arising difficulties in language comprehension are
hypothesized to stem from age-related working memory limitations
(Wingfield & Grossman, 2006), in turn linked to alterations in
prefrontal (Uekermann et al., 2008) and frontal cortices (Lee &
Federmeier, 2011), which play crucial roles in working memory and
inhibitory control. Conversely, the higher degree of preservation of
comprehension abilities compared to production might stem from
the higher burden imposed by the latter (Diaz et al., 2016), as well as
from difficulties to rely on contextual cues (Shafto & Tyler, 2014).
Figure 1 summarizes the attested language deficits and strengths in
the domains of production and comprehension in healthy ageing. In
the next section, we compare this pattern to the pattern of language
acquisition by a child, with the aim to test whether the former
inversely mimics the latter as anticipated by the hypothesis of
retrogenesis.
Does Language Decline in Ageing Inversely Mirror
Language Acquisition by the Child?
As observed in ageing, but also in conditions in which language is
disordered, child language acquisition exhibits a noteworthy vari-
ability with regard to onset times, rates, itineraries of development,
and ultimately, the pace of milestone achievement. Still, compared to
language changes during ageing, rather less studied, language acqui-
sition by the child has been researched much in depth. In general,
as summarized by Bates and Dick (2002), acquisition goes through
distinctive, although partially overlapping stages: (multi)word
comprehension (8–10 months), single word production (11–13
months), basic word combination (18–20 months), emergence of
first functional words (24–30 months), and use of simple syntactic
structures, including different types of sentences (i.e., declarative,
interrogative, imperative; 30–34 months). Throughout the whole
process, comprehension precedes production (Visser-Bochane et al.,
2020), although an inverse pattern can be observed with regard to
some specific aspects of language (e.g., word order or object
pronouns; see Hendriks, 2020). Concerning the two domains of
interest for our analysis (lexicon and syntax), word comprehension
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6BENÍTEZ-BURRACO AND IVANOVA

in children typically precedes expressive vocabulary acquisition
(Messer, 2015;Roemer et al., 2019), just as syntax understanding
precedes syntax production (Huttenlocher et al., 2002). During child
language development, vocabulary growth is a robust predictor of
grammar complexity (Dixon & Marchman, 2007). Earlier utterances
tend to be less syntactically complex and more lexically specific
(Theakston & Lieven, 2017). Lexical development does not reach
adult-level features and functionality, specifically automatization, up
until the age of 7–9, once the number of stored words enables to
create adult-like networks encompassing semantic and phonological
relations. Likewise, although more advanced syntax can be observed
in younger ages (Vasilyeva et al., 2008), longer sentences and more
diverse syntactic constructions are not typically found in children
below 5 years of age (Tolchinsky, 2004). Overall, child language
development progresses from simple structures to more complex and
interdependent constructions performing multiple functions.
Table 1 shows the parallels that exist between language itiner-
aries in children and elderly in the two domains of language under
scrutiny. It is worth noting that despite individual variation, it is
relatively easy to trace and describe the milestones of language
acquisition from the chronological standpoint, since language
development in the child builds on cognitive achievements with
robust biological basis. By contrast, associating age-related lan-
guage changes with specific lifetime stages is more complex. One
reason is that older adults show heterogeneous cognitive reserves
and different decline paces (see Kemper et al., 2001). Other
reasons are the differential effects of the sociolinguistic back-
ground, the education level, or even the type of tests used for
language assessment (Bastin et al., 2012;Stine-Morrow, 2007).
In our comparison, we have used the approximate landmarks
resulting from experimental studies that include chronological
references (i.e. Abrams & Farrell, 2010;Balota et al., 2002;
Barresi et al., 2000;Burke & Shafto, 2004;Caplan et al.,
2011;DeDe & Knilans Flax, 2016;Griffin & Spieler, 2006;
Kemper et al., 2001;McGinnis & Zelinski, 2003;Moscoso
Del Prado Martín, 2017;Ouyang et al., 2020;Pomareda et al.,
2019;Poulisse et al., 2019;Sung et al., 2017;aswellasother
works cited throughout the article).
In the remainder of this section, we provide a detailed comparison
of lexicon and syntax changes observed in childhood and in healthy
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Figure 1
Main Tendencies of Language Impairment and Preservation in Healthy Ageing
Note. Preserved and impaired language functions in healthy ageing in the domains of syntax (in pink) and lexical semantics (in green). The approximate axes
reflect general trends: the y-axis corresponds to the degree of preservation of a function (the higher the point, the better preserved the function), while the x-axis
sorts global linguistic functions (comprehension/production) subdivided into preserved and impaired spectra. Troughs of the lines are speculative. As noted,
healthy ageing accounts for difficulties in comprehension and production of complex sentences and sentences with embedded clauses (left and right), while
comprehension of simple and frequent syntactic structures and production of simple and coordinate clauses (middle) are preserved. Likewise, healthy ageing
accounts for impaired comprehension of nonliteral meanings and despecification of lexical competition and word-finding problems during production (left),
while global lexical comprehension and production (middle) are preserved. See the online article for the color version of this figure.
LANGUAGE RETROGENESIS AND LANGUAGE EVOLUTION 7

ageing (summarized in Figure 2). Further on, we discuss whether the
observed parallels and differences support the hypothesis of lan-
guage retrogenesis.
Deficits in Lexical Access During Production Due to
Phonological Disruption
As mentioned above, one of the first changes in healthy ageing
is associated with word-retrieval difficulties, which are caused
by constraints in matching phonological forms to concepts. During
language development, young children also experience word-finding
difficulties that originate from imprecise phonological representations
of words (Messer & Dockrell, 2006). Likewise, children from 8 to
10 years of age exhibit tip-of-the-tongue problems when trying to
access infrequent and lengthy words, as well as words from sparse
neighborhoods (Newman et al., 2018). In this domain, the patterns of
age-related impairment of lexical access are inverse to the patterns
followed by lexical access development in children.
Despecification and Literality in Word Comprehension
Children’s ability not to under- or overgeneralize word mean-
ings is acquired belatedly, usually after the age of five (Tolchinsky,
2004). Prior to that age, children frequently overextend words.
Indeed, younger children prefer semantic, rather than contextual
sources for interpreting the meaning of words and utterances
(Ryder & Leinonen, 2014). Understanding nonliteral meanings
is a late-acquired language ability. The same can be said of lexical
competition, which results from increasing lexical proficiency as
more and more words are being integrated into the lexicon
(Weighall et al., 2017). Thus, 7-year-old children only exhibit lexical
competition for highly familiar words. Later, lexical competition also
extends to words with lower neighborhood density (see Henderson
et al., 2013). At approximately age of 12, the lexicon becomes adult-
like, and word recognition results automatized (Weighall et al., 2017).
As discussed in the previous section, older adults tend to surface
lexical competition strategies by activating less predictable lexical
solutions. Overall, in this respect too, ageing seems to disrupt a
late-acquired language function.
Simplification of Syntactic Structures in Language
Production
As also mentioned above, lower memory capacities and increased
difficulties with lexical access result in syntactic simplification,
which is typically observed in older adults. Embedded utterances
and left-branching sentences become particularly challenging.
During language development, complex sentences become frequent
in the child’s discourse only after mastering embedding (5-year-olds),
juxtaposed sentences (3-year-olds) and two-word utterances (2-year-
olds; Visser-Bochane et al., 2020). Cognitive development is usually
regarded as a robust predictor of syntactic complexity, with the
number of clauses per sentence/utterance being a good marker of
later syntactic development (Nippold et al., 2005). In this respect,
early age-related difficulties with complex syntactic production
reverse their late acquisition in childhood.
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Table 1
Language Acquisition in Neurotypical Children and Language Impairment in Healthy Ageing: Chronological Stages and Associated
Phenomena
Main milestones of language acquisition by normotypical children Main language changes during healthy ageing
a
8–10 months Language comprehension onset Comprehension of frequent words Generally preserved until
very old age10–12 months Understanding first words
Comprehension of very simple (2–3-word)
grammatical structures
11–13 months Single words production Use of specialized vocabulary
18–20 months Short simple phrases production Production of noncomplex sentences
Production of simple utterances24–30 months First functional words appear during
production
30–34 months Production of different types of simple
sentences (declarative, imperative,
questions), and complete sentences
34 months Occasional production of complex sentences,
mainly compound
Frequent use of complex sentences
5 years+Production and comprehension of longer
sentences and larger repertoire of syntactic
constructions
Disruptions in the production and
comprehension of long sentences
74+years
First appearance of complex syntactic
utterances with embedded clauses
Problems with complex sentences with
embedded clauses and left-branching (focal)
utterances
[50–70]
b
70+years
7 years+Nonliteral meaning comprehension
Control for lexical competition, especially for
frequent words
Problems for understanding nonliteral
meanings
Despecification in lexical competition
70+years
8 years+Ongoing development for quick adjustment of
phonological form to low-frequent and long
words
Word-finding difficulties due to phonological
deadjustment to lexical form, mainly for
low-frequent, and semantically empty
words
[50–70]
b
70+years
a
We assume the extreme difficulty to assign language changes in ageing to any specific chronological milestones, mainly because of high intersubject
variability. This column is therefore inherently speculative, though based on real data originating from a selection of experimental studies.
b
The change
itself is long-lasting and progressive. The end age means the most disruptive point for language change.
8BENÍTEZ-BURRACO AND IVANOVA

Difficulties in Understanding Complex Syntactic
Structures
Simple syntactic structures (2–3-word utterances) are usually
understood by children very early (Koizumi et al., 2019). However,
complex sentence comprehension is more variable among children,
with success rates correlating with exposure levels to complex syntax
(Huttenlocher et al., 2002). As noted, it is only after the age of 5
that children usually achieve good command over complex syntax,
including the understanding of transformed structures, like interro-
gativesorpassives(Tolchinsky, 2004). Young children show a
preference for the semantic aspects of sentences. Accordingly,
when listening to embedded sentences describing two related events
(e.g., John started cooking dinner once he took a shower), they tend to
focus on the linear order in which the two clauses appear, instead of
focusing on the structural dependencies between the main clause and
the subordinate clause (Blything et al., 2015). This strategy can result
in comprehension problems (i.e., young children might infer that John
cooked first and had a shower only later). Likewise, they struggle to
correctly interpret two dependent clauses with different subjects (e.g.,
John saw the woman who works at the local library;Booth et al.,
2000). This way, syntax comprehension follows inverse developmen-
tal paths in children and elderly, at least with regard to noncanonical
constituent-order and complex sentences with dependent clauses.
Overall, if we compare the patterns of language development in
children and language changes in healthy ageing (see Figure 2), we
can conclude that ageing mainly affects late-acquired language
functions, thus supporting the hypothesis of language retrogenesis.
Before ending this section, we wish to mention two potential
limitations of this broad conclusion (which call for future research
in these domains). First, it is still an open question to which extent
changes in verbal behavior observed during healthy ageing involve
compensatory strategies, as found in patients with AD (e.g.,
Gutiérrez-Rexach & Schatz, 2016). Such strategies are particularly
noticeable in the discursive domain (Ivanova, 2020) and can partially
mask some of the language functions or structures compromised by
ageing. Second, we need to know whether this retrogenetic pattern, as
sketched above, is cross-linguistically common. Despite notable
individual differences, available evidence (which is still quite limited
in quantity and scope) points to a common track of ageing-related
changes in the neurocognitive substrate of language (see Peelle, 2019;
Shafto & Tyler, 2014, for an insightful overview of ageing-driven
changes in neurocognitive substrate for language). This finding would
support the view that language function declines similarly in speakers
of different languages. In their seminal study, Juncos-Rabadánand
Iglesias (1994) compared the performance of elderly speakers of 14
different languages, including Arabic, Hungarian, Norwegian, Bas-
que, or Bulgarian (to name but a few), and found a common pattern of
impairments across the entirse sample in spite of notable typological
language differences. More focused studies support this view in the
domains of phonological fluency (when comparing English, Spanish,
Danish, and Hebrew; Oberg & Ramírez, 2006), emotional vocabulary
(when comparing English and Spanish; Schrauf & Sanchez, 2004), and
overall verbal fluency and semantic associations (when comparing
English and Spanish; Rosselli et al., 2002). Most noticeable cross-
linguistic differences pertain to semantic-pragmatic phenomena, which
are more dependent on sociocultural constraints, particularly in the
domain of speech (see Gerstenberg et al., 2018, on differences between
German and French) and word categorization (see Rosselli et al., 2002,
on differences between English and Spanish).
In the last section of the article, we examine again the hypothesis
of language retrogenesis from the neurobiological perspective. As
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Figure 2
Age-Related Language Changes and Child Language Development Milestones
Note. Comparative overlap of language development in neurotypical children and language changes in healthy ageing. The figure illustrates
some major steps in the development of the child’s understanding abilities (in pink) and production abilities (in green). Colored bubbles suggest
inflexion points between late-developing language functions in children and early disrupted language abilities in healthy ageing. See the online
article for the color version of this figure.
LANGUAGE RETROGENESIS AND LANGUAGE EVOLUTION 9

mentioned in the previous sections, language changes during early
development boil down to cognitive specialization and, ultimately,
to brain development.
Language Retrogenesis and the Neurobiological
Substrate for Language
The above discussion about the parallels between language
acquisition by the child and language changes in healthy ageing
gives support for the hypothesis of language retrogenesis. Yet,
additional support can be expected from the consideration of
neurobiological evidence. Brain structure and brain (basic) func-
tions are shaped by biological factors, mostly genetic by nature.
Accordingly, they are expected to have stronger links to evolution-
ary concerns in the spirit of evo–devo theories in biology. Under the
retrogenetic view, brain regions/computations that are known to
develop/consolidate later during development are expected to be the
first to deteriorate/disrupt during ageing. In this section, we review
evidence supporting this possibility.
Ageing results in diverse structural and functional changes in the
brain. First, brain volume reductions are frequently observed. Vol-
ume reductions mainly affect frontal and medial temporal areas
(Wingfield & Grossman, 2006), particularly the lateral prefrontal
cortex and the hippocampus (Raz et al., 2004). In turn, volume
reductions are commonly associated with reduced synaptic density
and alterations in functional connectivity, including changes in
dendritic morphology, neuronal connectivity, and neurotransmitter
integrity (Wingfield & Grossman, 2006). Additionally, ageing-
driven changes in the gray matter of the cerebral cortex, specifically,
are frequently observed in ageing people. Reductions in gray matter
density are commonly found in the temporal lobe (Fletcher et al.,
2018) and in the prefrontal cortex (Spreng & Turner, 2019), but also
in the frontal lobes, which are known to mature later in life (Rathi
et al., 2014). Third, neuroanatomical studies also report changes in
the white matter as the brain ages (Nilsson et al., 2014). Myelination
is a structural hallmark of consolidated functions. The disruption of
brain myelination patterns starts from the least myelinated brain
regions, which are the last to develop (Reisberg et al., 1999). Finally,
ageing also entails specific molecular changes affecting brain func-
tion, including changes in gene methylation patterns (Harman &
Martín, 2020) or dopaminergic transmission (Manenti et al., 2013).
All the brain areas highlighted above are crucially involved in
language processing and their disrupted connectivity accounts for
language problems in diverse clinical conditions. For example, the
prefrontal cortex supports working memory abilities (Bahmani
et al., 2019;Dienel et al., 2022;Funahashi, 2017;Wischnewski
et al., 2021), which are necessary for computing complex syntactic
structures, among other aspects of grammar. Likewise, alterations of
frontotemporal connectivity account for the difficulties with word
encoding experienced by people with schizophrenia (Wolf et al.,
2007). To mention one more case, decreased white matter density in
a left-sided frontotemporal network has been found in children with
developmental language disorder (Jäncke et al., 2007).
It is not easy to establish direct comparisons between changes in the
ageing brain, of the sort discussed above, and the structural and
functional changes experienced by the child brain during language
acquisition. One important reason is thatthe latter are more influenced
by genetic factors, while the former are more dependent on age-driven
physiological deterioration. But it is also true that ageing also depends
on genetic factors (Revelas et al., 2018;Walter et al., 2011), whereas
language acquisition depends as well on external stimuli. Conse-
quently, some reliable comparisons between both processes can be
established. Below, we briefly review evidence supporting the view
that the development of the language networks also mirrors, though
inversely, the deterioration of these networks with ageing.
Overall, language-related areas exhibit protracted development
over the life span (Pujol et al., 2006). Initially, language processing
seems to depend on functionally unspecific areas with a more
dispersed localization (Messer, 2015). Likewise, young children
show less left lateralization for language (Lidzba et al., 2011), with
lateralization to the left occurring with age, as the left hemisphere
areas associated with language become progressively active
(Szaflarski et al., 2006; see Figure 3A, for details). To an important
extent, these modifications parallel, though inversely, the changes
that can be observed in the ageing brain. First, the ageing brain
shows a noteworthy neurofunctional dispersion that stems from
large-scale reorganizations impacting on diverse cognitive abilities
(Bagarinao et al., 2019). Furthermore, some structural changes in the
ageing brain follow an anterior–posterior gradient, affecting late-
myelinated language-supporting areas in the first instance, like the
left inferior frontal gyrus (Diaz et al., 2016). A parallel decline along
a superior–inferior gradient can be observed too, impacting as well
on white matter tracts important for language processing, specifi-
cally the arcuate fasciculus (more on this track below), and ulti-
mately, on speech production and syntax (Diaz et al., 2016). In
general, in both healthy and pathological ageing, more pronounced
changes are observed in late-myelinated brain areas (Gao et al.,
2011), which seemingly accounts for the (pattern and timing of)
slowdown or breakdown in the cognitive functions they support
(Brickman et al., 2012; see Figure 3B, for details). Finally, healthy
ageing also results in a bilateral activation of language areas, which
contrasts with the left-lateralized patterns observed at younger ages
(see Hommet et al., 2008, among others), but which resembles what
can be observed in young children (see Plessen et al., 2014, and
Zhou et al., 2013, for data presentation and discussion).
Overall, the structural and functional reorganization of the brain
during ageing has been characterized as a process of dedifferentia-
tion or compensation (see Wingfield & Grossman, 2006,fora
review). From that perspective, ageing conveys a progressive
reduction of cortical specialization for language that results in
the consequent recruitment of other brain areas for language-
related tasks. For instance, in order to achieve good understanding
of complex sentences, older adults involve left dorsal inferior
frontal and right temporal–parietal regions (involved in working
memory and syntax processing) to compensate for the ageing-
related underactivation of left temporal–parietal areas (which
are part of the short-term auditory–phonological buffer that
enables information maintenance during sentence processing;
Wingfield & Grossman, 2006). Furthermore, when this compen-
satory strategy fails, ageing speakers can also activate dorsolateral
prefrontal cortex areas, which are in charge of general-purpose
problem-solving tasks (Wingfield & Grossman, 2006). Interest-
ingly for the retrogenetic view, children over 10 exhibit increased
activations of the left temporal–parietal junction and the right
superior temporal gyrus when processing complex sentences
(Yeatman et al., 2010). This way, ageing can be said to result
in a progressive dedifferentiation and despecialization of brain
activation patterns during language processing. As noted, most of
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10 BENÍTEZ-BURRACO AND IVANOVA

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Figure 3
Graphical Comparison of the Developing and the Ageing Brains
Note. A. Lateralization patterns for language in child brain and ageing brain. Progressive lateralization and reorganization patterns for language in a child
brain (with a focus on Age 5) and an ageing brain, both assumed for right-handedness as more dominant pattern. In Image 3A.1 (left), the darker color indicates
later and/or protracted lateralization in children. In a child brain, language lateralization to the left boosts around Age 5, specifically in Broca’s area (left IFG,
BA44 and BA45) and angular gyrus (BA39; Szaflarski et al., 2006). Lateralization of Wernicke’s area (BA21 and BA22, also BA39) is the mostly pronounced,
yet not dominant until Age 7 (Berl et al., 2014). Before pronounced left lateralization, children recruit right homologs of language basic network (mainly IF and
ST cortices), including Broca’s and Wernicke’s areas; involvement of middle frontal gyrus (BA46) remains stable, with frequent bilateralization. Crucially,
Broca’s area specialization is more protracted developmentally. In Image 3A.2 (right), the darker color indicates more preserved specialization of lateralization
in healthy ageing. In ageing brain (right), atrophy of language-related areas coincides with recruitment of several areas in the right hemisphere, mainly
dorsolateral prefrontal cortex (BA10 and BA46) and temporoparietal cortex. Ageing speakers show less activation of several language-supporting areas, like
left inferior prefrontal cortex (BA45 and BA47) or posterolateral temporal-parietal cortex (BA39), but instead they tend to recruit additional, non-language-
supporting brain areas, for example, temporal left pole-inferior temporal gyrus (BA20 and BA38), posterior cingulate cortex (BA23 and BA31) or anterior
temporal lobe (BA38) (Antonenko et al., 2013;Lacombe et al., 2015;Pistono et al., 2021). Images adapted from Pixabay (http://www.pixabay.com).
B. Myelination patterns of language-supporting areas in healthy mature brain, child brain, and ageing brain. In a healthy mature brain (left), the most prominent
language-supporting areas, according to Sakai (2005) and Ardila et al. (2016), are colored. Broca’s complex includes the left inferior frontal gyrus (F3T/F3O),
BA45 and BA47; the left inferior frontal gyrus, specifically opercular pars and triangular pars, BA44 and BA45, and the left middle frontal gyrus, BA46.
Wernicke’s complex includes the posterior superior temporal gyrus, BA22; the middle temporal gyrus, B21; the auditory cortex, BA41 and BA42. Extended
Wernicke’s area also includes BA20, BA37, BA38, BA39 and BA40. FA (the arcuate fasciculus) is considered to be the most important white matter bundle for
language network in the brain. In the figure depicting the language-supporting areas in a healthy child brain (center), the late-myelinated areas are highlighted in
darker colors. These include some parts of the Broca’s complex areas, related to language production and complex language support, as well as the FA,
responsible for integration–production networking. In the figure of the healthy ageing brain (right), the darker areas correspond to the language-supporting areas
that are more susceptible to demyelination. These include the left IFG and the FA. Importantly too, impairments in syntactic and semantic processing in ageing,
to which the IFG contributes significantly, are hypothesized to be related to the reinforcement of reactive aggression during ageing, giving additional support to
the view that language-related areas mediate executive abilities involved in the cortical regulation of both aggression and combination, including syntax (Miller
et al., 2008). IFG =inferior frontal gyrus; BA =Brodmann area; IF =inferior frontal (cortex); ST =superior temporal (cortex); FA =arcuate fasciculus. Images
adapted from Pixabay (https://www.pixabay.com). See the online article for the color version of this figure.
LANGUAGE RETROGENESIS AND LANGUAGE EVOLUTION 11

these changes recapitulate, in a reverse way, the changes that
happen in the brain of children as they grow.
Language Retrogenesis and Language Evolution
As discussed in the previous sections, available linguistic, cogni-
tive, and neurobiological evidence support to a great extent the
hypothesis of language retrogenesis, specifically with regard to the
lexicon and syntax. In this section, we will address the links between
this retrogenetic view and the way in which language evolved in the
species. As we lack direct evidence of prehistoric language(s), and
because paleoneurological, but also paleoanthropological and pa-
leogenetic evidence is becoming increasingly available, we find it
safer to move the discussion to the domain of the biological substrate
for language.
In general, the developmental path of the neural substrate for
language parallels the evolution of the components of the
language-ready brain. For instance, Friederici (2012) shows that the
dorsal pathway of the language network, which is involved in
processing of hierarchical structures, is not myelinated in the newborn,
and only becomes fully functional after maturation. This pathway is
regarded by many, for example Aboitiz et al. (2010), as a human-
specific innovation. Likewise, the arcuate fasciculus, a nervous bundle
supporting complex syntax processing, is underdeveloped in children,
particularly, in subjects with syntactic processing problems (Friederici
& Gierhan, 2013). The integrity of the left arcuate fasciculus (or, more
generally, the language network to which this fasciculus contributes)
declines with age (Ikuta et al., 2020). Importantly, this decline
correlates with diverse age-related language changes, including the
tip-of-the-tongue phenomena (Shafto & Tyler, 2014)ordifficulties
with syntactic comprehension (Antonenko et al., 2013). The expansion
of the arcuate fasciculus is regarded as a major milestone in language
evolution in humans (Rilling, 2014). Among other things, this track is
less developed, or even absent, in nonhuman primates (Rilling et al.,
2008). To give a final example, the left inferior frontal gyrus is one of
the late-myelinated cortical areas that are the first to be affected by
ageing (Diaz et al., 2016). This impairment seemingly contributes to
the notable difficulties exhibited by aged people with passive sen-
tences (Sung et al., 2017), as the bilateral inferior frontal gyrus is
responsible for thematic role assignment (Mack et al., 2013). From an
evolutionary perspective, when compared to macaques, the human left
inferior frontal gyrus shows increased functional integration and
coupling, as well as increased myelination, which are reminiscent
of the changes experienced by the child brain as it develops (Wang
et al., 2020;seeFigure 3B for more details). Not surprisingly
perhaps, passives are regarded a late innovation of natural lan-
guages, as suggested by mathematical modeling of early steps in
the evolution of human language (Nowak & Krakauer, 1999).
Overall, this evidence supports the view that the areas that are
late-myelinated in childhood, and which are also the earliest to be
disrupted in ageing, are responsible for some of the most computa-
tionally demanding aspects of language, and are also the most recent
to have evolved in the species. As we advanced in Section Language
Retrogenesis and “Degraded”Form of Language above, the primary
reason for these parallels is that recently evolved networks are
endowed with weaker protection mechanisms to damage, either
ontogenetic or acquired, because of their evolutionary novelty
(cf. Toro et al., 2010 or Pattabiraman et al., 2020, for discussion).
As a consequence, they become deteriorated first during ageing, and
are the last to be developed during childhood.
Conclusions and Prospects
The main conclusion of our analysis is that language acquisition in
children and language change/loss in healthy ageing generally align
with the principle of retrogenesis. This is particularly true for complex
language tasks, such as comprehension of constrained lexicon, or
comprehension and production of complex syntax. Still, certain
mismatches can be observed in this inverted mirror pattern, which
are seemingly justified by the richer experience in language use
exhibited by aged speakers. Accordingly, while elderly people can
still, although less efficiently, rely on the background knowledge to
compensate for language difficulties (see Federmeier et al., 2010),
children only gradually develop the needed abilities to enrich linguistic
meanings relying on relevant contextual cues (Ryder & Leinonen,
2014). Furthermore, because of the longer use, both declarative and
procedural aspects of language are better trained in the elderly and,
thus, are more consistent and resistant to ageing. Finally, social
networking affects language structures and use differently in both
groups. That said, we have also found neurobiological and evolution-
ary evidence supporting the feasibility of language retrogenesis. In
fact, language retrogenesis can be plausibly explained by the way in
which language has evolved, with aspects of language introduced
more recently being more (and earlier) affected in ageing and emerg-
ing later in children, and vice versa. Seemingly this explains as well
why the less structurally complex and less culturally conditioned
components of the language follow this retrogenetic model to a lesser
extent. This is particularly true for phonology (recall our detailed
discussion in Section Language and Cognitive Changes in Healthy
Ageing above). This pattern is also coherent with some linguistic
views, for example, Jackendoff’s (1999) or Progovac’s (2015),ac-
cording to which the earlier steps in the evolution of grammar are not
subsequently replaced, but, rather, elaborated upon (this also being
true for language ontogeny, we wish to add).
Hypotheses of this kind need to be properly tested. More compar-
ative studies are clearly needed to provide additional support to (and,
also, to refine) the hypothesis of retrogenesis, particularly with focus
on the aspects of language we highlight in the article and with direct
comparisons between children and aged people. We also need more
data from non-WEIRD (Western, Educated, Industrialized, Rich, and
Democratic) societies, as cultural and sociological environments in
which people grow and age can differ significantly, with differences
having an impact on language acquisition and loss (although for the
hypothesis of retrogenesis, we are interested in the commonalities of
these phenomena across different populations). A growing body of
work urges to include linguistic typology as a key factor in studies on
human neurocognition (Calvo et al., 2018;Kemmerer, 2014). Like-
wise, we need to continue delving into the psycholinguistic aspects of
language acquisition by the child and language variation in the
elderly, as this approach can provide a clearer picture of their
cognitive underpinnings. Furthermore, it will also facilitate a more
accurate neurobiological account of language acquisition and loss, of
the sort we have sketched in the article. Genetic studies could help us
learn more about the biological causes of the observed variation when
acquiring or losing a language. In turn, better neurobiological and
genetic characterizations of language in ageing are the key to conduct
comparative studies in other species, aimed to refine our knowledge
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12 BENÍTEZ-BURRACO AND IVANOVA

of the evolutionary trajectory of the language-related areas that are
impacted differently by ageing. Eventually, studies using ancient
DNA might result in more detailed accounts of these evolutionary
changes, particularly, of gene modifications influencing language
changes in ageing, because of the deep connection between evolution
and development, of the sort we have stressed so far.
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Received December 4, 2021
Revision received October 18, 2022
Accepted November 8, 2022 ▪
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18 BENÍTEZ-BURRACO AND IVANOVA