Independent Representations of Verbs and Actions - Am MIT spezialisierte KI-Forschung

Independent Representations of Verbs and Actions
in Left Lateral Temporal Cortex

Marius V. Peelen1,2, Domenica Romagno3, and Alfonso Caramazza1,2

Abstrakt

■ Verbs and nouns differ not only on formal linguistic grounds
but also in what they typically refer to: Verbs typically refer to ac-
tionen, whereas nouns typically refer to objects. Prior neuroimaging
studies have revealed that regions in the left lateral temporal cor-
tex (LTC), including the left posterior middle temporal gyrus
(pMTG), respond selectively to action verbs relative to object
nouns. Other studies have implicated the left pMTG in action
Wissen, raising the possibility that verb selectivity in LTC
may primarily reflect action-specific semantic features. Hier, verwenden
functional neuroimaging, we test this hypothesis. Participants per-
formed a simple memory task on visually presented verbs and
nouns that described either events (z.B., “he eats” and “the
conversation”) or states (z.B., “he exists” and “the value”). Verb-

selective regions in the left pMTG and the left STS were defined
in individual participants by an independent localizer contrast
between action verbs and object nouns. Both regions showed
equally strong selectivity for event and state verbs relative to se-
mantically matched nouns. The left STS responded more to states
than events, whereas there was no difference between states and
events in the left pMTG. Endlich, whole-brain group analysis re-
vealed that action verbs, relative to state verbs, activated a cluster
in pMTG that was located posterior to the verb-selective pMTG
clusters. Zusammen, these results indicate that verb selectivity in
LTC is independent of action representations. We consider other
differences between verbs and nouns that may underlie verb
selectivity in LTC, including the verb property of predication. ■

EINFÜHRUNG

Nouns and verbs are elemental grammatical units of
most, if not all, human languages. Nouns and verbs are
identified on the basis of formal morphosyntactic features.
Zum Beispiel, nouns bear nominal markers, and verbs bear
verbal markers. Darüber hinaus, verbs and nouns are associated
with distinct syntactic patterns and play different roles in
Sätze. The categories of verb and noun are, daher,
formally discrete, in so far as they are encoded by clear-cut
linguistic units.

Jedoch, nouns and verbs differ not only on formal
linguistic grounds but also differ in what they typically
refer to. Nouns typically (but not always) refer to entities,
whereas verbs typically (but not always) refer to events. Es
is possible to identify prototypical members of each cate-
gory. Prototypical verbs denote agentive dynamic actions,
such as to walk and to build; prototypical nouns denote
concrete three-dimensional individualized entities, solch
as the table and the professor. But verbs may also refer to
Staaten (to stay) or relations (to belong), and nouns may also
refer to events (the wedding) or conditions (the tempera-
tur; Hopper & Thompson, 1984; Lyons, 1977).

Prior functional neuroimaging studies have revealed
brain regions selectively engaged in verb processing (Bedny,
Caramazza, Pascual-Leone, & Sachsen, 2012; Willms et al.,
2011; Bedny, Caramazza, Grossman, Pascual-Leone, &

1University of Trento, 2Harvard Universität, 3University of Pisa

Sachsen, 2008; Shapiro, Moo, & Caramazza, 2006; Yokoyama
et al., 2006; Kable, Kann, Wilson, Thompson-Schill, &
Chatterjee, 2005; Perani et al., 1999), although there ap-
pears to be considerable variability across studies in the
specific regions reported (Crepaldi, Berlingeri, Paulesu, &
Luzzatti, 2011), which is presumably related to differences
in tasks and/or stimuli. The most consistent finding across
studies is a selective involvement of the left lateral temporal
Kortex (LTC), including the posterior middle temporal
gyrus (pMTG), in verb processing (Bedny et al., 2008,
2012; Crepaldi et al., 2011; Willms et al., 2011; Burton,
Krebs-Noble, Gullapalli, & Berndt, 2009; Tyler, Randall, &
Stamatakis, 2008; Shapiro et al., 2006; Perani et al., 1999).
Because these prior studies have generally contrasted
action verbs with object nouns, it is unknown whether
verb-selective responses in LTC reflect lexical, syntactic,
and/or semantic differences between verbs and nouns.

Interessant, the left LTC has also been consistently
implicated in conceptual action and tool knowledge
(Kemmerer, Rudrauf, Manzel, & Tranel, 2012; Campanella,
DʼAgostini, Skrap, & Shallice, 2010; Kalenine, Buxbaum, &
Coslett, 2010; Martin, 2007; Kable et al., 2005; Tranel,
Kemmerer, Adolphs, Damasio, & Damasio, 2003; Chao,
Haxby, & Martin, 1999; Martin, Haxby, Lalonde, Wiggs, &
Ungerleider, 1995). This raises the possibility that verb se-
lectivity in the left LTC may be driven primarily by semantic
differences between action verbs and object nouns.
In der Tat, several studies have found that sentences referring
to actions activate regions in LTC (Wallentin et al., 2011;

Zeitschrift für kognitive Neurowissenschaften 24:10, S. 2096–2107

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Deen & McCarthy, 2010; Rueschemeyer, Glenberg, Kaschak,
Mueller, & Friederici, 2010; Saygin, McCullough, Alac, &
Emmorey, 2010; Hauk, Davis, Kherif, & Pulvermuller,
2008; Wallentin, Lund, Ostergaard, Ostergaard, &
Roepstorff, 2005; Davis, Meunier, & Marslen-Wilson,
2004), including a verb-selective region in STS (Davis
et al., 2004). Jedoch, a recent study investigating re-
sponses in functionally localized verb-selective LTC regions
showed that these regions are not modulated by the
amount of visual motion or motor activity associated with
verbs, with equally selective responses to verbs such as
to jump and to think (Bedny et al., 2008). Außerdem,
verb-selective LTC responses have also been reported in
congenitally blind participants (Bedny et al., 2012), exclud-
ing the possibility that selectivity for action verbs in LTC
relates to visual motion.

Wichtig, Jedoch, actions also differ from objects at
a more abstract conceptual level, such as the understand-
ing of actions (but not objects) as dynamic events un-
folding over time. Daher, rather than or in addition to
representing basic sensory motor features, verb-selective
LTC may very well represent these more abstract semantic
differences between actions (including mental actions) Und
Objekte. In this study, we tested this hypothesis by dis-
entangling grammatical category from prototypical seman-
tic features of verbs and nouns.

If verb selectivity in LTC reflects the retrieval of event
concepts, we would expect (1) that activity in LTC would
be relatively low to verbs that do not refer to events (z.B.,
to include) relative to verbs that do (z.B., to talk) Und
(2) that activity in LTC would be relatively high to nouns
that refer to events (z.B., the run) relative to nouns that
do not (z.B., the identity). To test these predictions, Wir
performed two experiments in which grammatical cat-
egory (verb, noun) and dynamicity (state, Ereignis) war
manipulated in a 2 × 2 factorial design. Both experiments
also included separate action verb and object noun con-
ditions to localize LTC regions reported in previous stud-
ies (Bedny et al., 2008, 2012; Willms et al., 2011; Burton
et al., 2009; Tyler et al., 2008; Shapiro et al., 2006; Perani
et al., 1999). To ensure that results were not specific to
a particular stimulus set, the two otherwise identical
experiments differed in the stimulus material used: In
Experiment 1, all the noun phrases had a shared root
with a verb (z.B., the desire), whereas in Experiment 2
none of the noun phrases had a shared root with a verb.
Außerdem, in Experiment 1 all verb phrases had an
animate subject (“he” or “she”), whereas in Experiment 2
all verb phrases had an inanimate subject (“this”).

METHODEN

Teilnehmer

A total of 27 healthy adult volunteers (12 Frauen; mean
age = 26 Jahre, age range = 20–36 years) participated in
the study (Experiment 1: n = 15; Experiment 2: n = 12).

All participants were native Italian speakers and right-
handed and had normal or corrected-to-normal vision
and no history of neurological or psychiatric disease. Par-
ticipants gave written informed consent for participation
in the study, which was approved by the human research
ethics committee of the University of Trento.

Two participants (one in each experiment) were ex-
cluded due to excessive head motion (>4 mm in any di-
rection across the experiment). One participant (aus
Experiment 1) was excluded due to low performance
(mean accuracy of <3 standard deviations of the group mean). This left a total 13 participants in Experiment 1 and 11 2. Stimuli For both experiments, stimuli consisted 20 short phrases per condition (in Italian), such as “she talks” and “the temperature” (Appendix). There were four experi- mental conditions: event verbs (EV; e.g., “this arrives”), event nouns (EN; “the conference”), state (SV; e.g., contains”), and (SN; iden- tity”). Two additional conditions included to localize regions previously implicated verb processing: action verbs (AV; “he jumps”) object (ON; e.g., “the lamp”). We use label “action verbs” (rather than “event verbs”) for localizer because all phrases denoted motor actions. It should be noted, how- ever, that there is considerable semantic overlap between the action all verbs describe events, many describe motor actions (see Appendix). The noun conditions purpose having a statistically independent selective ROIs, one used similar those previous studies. Event dynamic activities unfolding over time, chases” arrives,” whereas state verbs states or represented above time, exists” includes.” Event and state paralleled semantics, they denote dynamic like destruction” excursion,” or conditions, existence” “the temperature.” phrases experimental con- ditions matched cumulative form frequency (Bertinetto et al., 2005), word length letters, gram- matical gender pronouns preceded verbs. one-argument verbs, as to walk, two-argument build. The number arguments was across conditions. Different two experiments (Appendix). AV ON localize ROIs identical experiments. For Experi- ment 1, EV SV had an animate subject (“he” or “she”), 2, phrases had inanimate (“this”). In all Peelen, Romagno, Caramazza 2097 D o w n l o a d e d f r o m l l >297 mm3) was less than 5% ( P < .05). This threshold was adopted unless otherwise specified. Definition of ROIs ROIs were defined in individual participants and were re- stricted to voxels that were significantly activated in the corresponding group analysis (the results of the group analyses are given in Table 1). The same contrast that was used in the group analysis (AV > ON) was also used
in each individual participant, applying an individual par-
ticipant voxel threshold of p < .05 (uncorrected). No cluster size threshold was applied at the individual partic- ipant level. The first ROI, pMTG, defined by the contrast AV > ON, could be localized in all 24 participants and had

2098

Zeitschrift für kognitive Neurowissenschaften

Volumen 24, Nummer 10

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effects group analysis. Wie erwartet, significant activity was
observed in the left LTC (Tisch 1). The activity in LTC
consisted of a cluster in the pMTG and a more anterior
cluster in the MTG/STS. Verb-selective activity was also
observed in the left TPJ and the right inferior occipital
gyrus. A region in the left medial fusiform gyrus responded
more to object nouns than to action verbs (Tisch 1).

To test whether verb-selective regions could also be ob-
served for verbs that do not refer to actions or events, Wir
contrasted state verbs with state nouns. This contrast again
yielded significant activity in left LTC (Tisch 1). No regions
showed more activity to nouns than verbs in this analysis.
What is the relation between action and verb represen-
tations in LTC? To address this question, we compared
LTC activity to three different contrasts at more lenient
statistical thresholds ( P < .005, uncorrected), limiting the analysis to the LTC. The first contrast was between action verbs and object nouns, a contrast expected to ac- tivate both action and verb representations. The second contrast, between state verbs and state nouns, probes verb but not action representations. The third contrast, between action verbs and state verbs, probes action but not verb representations. The results are displayed in Figure 1. Interestingly, an action-selective cluster (Figure 1; AV > SV) was found posterior to verb-selective clusters
(Figur 1; SV > SN), indicating that nearby but distinct
clusters may represent verbs and actions. At this relatively
lenient threshold, the contrast between action verbs and
object nouns activated both verb- and action-selective
cluster (Figur 1; AV > ON). At more stringent thresholds
(Tisch 1), the AV > ON contrast primarily activated verb-
selective regions.

Endlich, we tested for differences between responses to
events and states, averaged across grammatical category.
No regions responded differentially to these categories at

Tisch 1. Overview of Activated Regions in Whole-brain
Random-effects Group Analyses (n = 24), Thresholded at
P < .05 (Corrected for Multiple Comparisons) Coordinates Contrast Region x y AV > ON

Left MTG/STS −51 −30

Left pMTG

Left TPJ

Right IOG

−49 −53

−57 −46

32 −80

−5

ON > AV

Left FG

−26 −33 −17

SV > SN

Left MTG/STS −61 −25

Volumen
(mm3)

567

1917

324

1944

864

405

Coordinates are of the center of mass of the clusters, in Talairach space.

IOG, inferior occipital gyrus; FG, fusiform gyrus.

a mean size of 893 mm3. The second ROI, MTG/STS,
defined by the contrast AV > ON, could be localized in
19 Teilnehmer (9 of Experiment 1, 10 of Experiment 2),
and had a mean size of 347 mm3.

ERGEBNISSE

Verhaltensergebnisse

Mean accuracy did not differ between AV and ON condi-
tionen (AV: 96.5%; ON: 96.9%; T(23) = −0.3, p = .74). Mean
RT did not significantly differ either (AV: 834 ms; ON:
819 ms; T(23) = 1.1, p = .29).

Accuracy and RT of the experimental conditions (EV,
EN, SV, SN) were analyzed in 2 × 2 ANOVAs with Dynam-
icity (Ereignis, state), and Grammatical Class (verb, noun) als
factors. There was a main effect of Grammatical Class, F(1,
22) = 5.3, p = .032, mit (slightly) higher accuracy for
nouns (97.8%) than verbs (96.4%). There was no significant
effect of Dynamicity, F(1, 22) = 2.7, p = .11, indicating
equally high accuracy for events (96.5%) and states
(97.7%). The interaction between Grammatical Class and
Dynamicity was not significant ( p = .12). For RT, Dort
was a main effect of Grammatical Class (F(1, 22) = 5.1,
p = .034), with faster responses to nouns (813 ms) als
to verbs (831 ms). There was again no significant effect
of Dynamicity, F(1, 22) = 0.6, p = .45, indicating equally
fast responses to events (819 ms) and states (825 ms).
The interaction between Grammatical Class and Dynam-
icity was not significant ( p = .78).

Results of Whole-brain Group Analyses

The first analysis was aimed at replicating and localizing
verb-selective responses in LTC. Following earlier work
(Bedny et al., 2008, 2012; Willms et al., 2011; Tyler et al.,
2008; Shapiro et al., 2006), we contrasted activity in re-
sponse to action verbs (z.B., “he jumps”) with activity to
object nouns (z.B., “the lamp”) in a whole-brain random-

Figur 1. Verb- and action-selective clusters in the left LTC. Ergebnisse
of three independent contrasts are shown at p < .005 (uncorrected): action verbs versus object nouns (AV > ON; top row), state verbs versus
state nouns (SV > SN; middle row), and action verbs versus state verbs
(AV > SV; bottom row). These results reveal two verb-selective LTC
clusters, in pMTG and MTG/STS, for nonaction verbs (SV > SN) Und
also indicate that distinct regions encode verbs (SV > SN) and actions
(AV > SV).

Peeling, Romagno, and Caramazza

2099

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the corrected statistical threshold. At a more lenient
threshold ( P < .005, uncorrected), activity was observed in the left LTC (xyz = −47, −30, −2), with stronger re- sponses to states than events. Results of ROI Analysis To test whether verb-selective responses in left LTC re- flect the retrieval of event concepts, we compared re- sponses to the four experimental conditions (EV, EN, SV, SN) within LTC ROIs defined by the independent contrast between AV and ON. In the whole-brain group analysis (at p < .05, corrected), this contrast revealed activ- ity in two separate regions of LTC (pMTG and MTG/STS; Table 1; note that these regions merged at the more le- nient threshold displayed in Figure 1). Responses in both these regions were further investigated by localizing them in individual participants (see Methods). Responses were analyzed for each ROI separately using mixed 2 × 2 × 2 ANOVAs with Experiment (1, 2), Dynamicity (event, state), and Grammatical Class (verb, noun) as factors. pMTG, localized in individual participants, showed a main effect of Grammatical Class, F(1, 22) = 29.3, p = .00002, with stronger responses to verbs than nouns (Figure 2). This result indicates that verb selectivity in pMTG generalizes to semantically matched verb–noun contrasts. The interaction between Grammatical Class and Experiment approached significance, F(1, 22) = 4.2, p = .054, with a stronger difference between verbs and nouns in Experiment 2, t(10) = 6.6, p = .00006, than in Experiment 1, t(12) = 2.2, p = .052. There were no sig- nificant effects involving the factor Dynamicity (main effect: F(1, 22) = 0.0, p = .95; Dynamicity × Grammatical Class: F(1, 22) = 2.7, p = .12; Dynamicity × Experiment: F(1, 22) = 1.3, p = .28; Dynamicity × Experiment × Grammatical Class: F(1, 22) = 0.0, p = .91). A similar pattern of results was obtained when pMTG was localized based on group-average activity without individual participant voxel selection: a main effect of Grammatical Class (F(1, 22) = 19.5, p = .0002), an inter- action between Grammatical Class and Experiment (F(1, Figure 2. Responses to the four experimental conditions in left LTC ROIs (pMTG and MTG/STS), defined by the contrast AV > ON. Error
bars indicate within-subject SEM.

22) = 5.9, p = .024), and again no significant effects in-
volving the factor Dynamicity ( p > .18, for all tests).

The MTG/STS ROI, defined by the contrast AV > ON in
individual participants, showed a main effect of Gramma-
tical Class, F(1, 17) = 25.6, p = .0001, with stronger re-
sponses to verbs than nouns (Figur 2). The interaction
between Grammatical Class and Experiment was signifi-
kippen, F(1, 17) = 5.9, p = .027, with a stronger difference
between verbs and nouns in Experiment 2, T(9) = 4.7, p =
.001, than in Experiment 1, T(8) = 2.3, p = .052. Dort
was a significant main effect of Dynamicity, F(1, 17) =
8.5, p = .010, with stronger responses to states than
Veranstaltungen (Figur 2). There were no significant interactions
between Dynamicity and the other factors (Dynamicity ×
Grammatical Class: F(1, 17) = 2.8, p = .11; Dynamicity ×
Experiment: F(1, 17) = 2.3, p = .14; Dynamicity ×
Experiment × Grammatical Class: F(1, 17) = 0.0, p = .97).
A similar pattern of results was obtained when MTG/
STS was localized based on group-average activity without
individual participant voxel selection: a main effect of
Grammatical Class, F(1, 22) = 16.2, p = .0006; an inter-
action between Grammatical Class and Experiment, F(1,
22) = 4.8, p = .039; and again a significant main effect
of Dynamicity, F(1, 22) = 5.0, p = .035, with stronger
responses to states than events.

Endlich, we tested whether verb-selective regions in
LTC distinguished between the three verb types (Aktion,
Ereignis, state) and/or between the three noun types (Objekt,
Ereignis, state). Because the ROIs used in the previous anal-
yses were defined by the localizer contrast between action
verbs and object nouns, comparing verb/noun types within
these ROIs would be statistically biased toward action verbs
and against object nouns. daher, we defined verb-
selective regions (pMTG and MTG/STS) using the contrast
between all verbs and all nouns based on group average
Aktivität. Within these regions, we then tested for differ-
ences between the verb and noun types in 2 × 3 ANOVAs,
with Experiment (1, 2) and Verb or Noun type (Veranstaltungen,
Staaten, actions/objects) as factors.

pMTG showed no main effect of Verb type, F(2, 21) =
2.4, p = .11, and no interaction between Verb type and
Experiment, F(2, 21) = 0.8, p = .45. MTG/STS showed a
main effect of Verb type, F(2, 21) = 4.3, p = .028, but no
interaction between Verb type and Experiment, F(2, 21) =
1.6, p = .22. The main effect of Verb type was driven by
a stronger response to state verbs than to event verbs,
T(23) = 2.9, p = .009, confirming the results of the anal-
yses in MTG/STS defined by AV > ON. There were no
significant differences between action and event verbs,
T(23) = 1.7, p = .11, or between state and action verbs,
T(23) = 1.3, p = .20.

pMTG showed a main effect of Noun type, F(2, 21) =
5.7, p = .011, but no interaction between Noun type and
Experiment, F(2, 21) = 1.1, p = .34. The main effect of
Noun type was driven by a lower response to object nouns
relative to both state and event nouns, T(23) > 2.3, P < .05, for both tests. There was no significant difference between 2100 Journal of Cognitive Neuroscience Volume 24, Number 10 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f t . / u s e r o n 1 7 M a y 2 0 2 1 state and event nouns, t(23) = 1.4, p = .18. The main effect of Noun type in MTG/STS did not reach significance, F(2, 21) = 2.9, p = .078, and there was no interaction between Noun type and Experiment, F(2, 21) = 1.7, p = .20. Pairwise t tests showed a stronger response to state nouns than ob- ject nouns, t(23) = 2.2, p = .039, but no difference between state nouns and event nouns, t(23) = 0.7, p = .52, or be- tween event nouns and object nouns, t(23) = 1.3, p = .20. DISCUSSION This study tested whether and how neural selectivity for verbs in the left LTC is related to the representation of con- ceptual action knowledge. We first replicated previous studies by contrasting action verbs with object nouns. This contrast gave significant verb-selective activity in a region in the left MTG/STS and a more posterior region in the left pMTG that was present in all 24 participants. Verb-selective activity in LTC was also observed for the contrast between nonaction (state) verbs and state nouns, with activity in both pMTG and MTG/STS at more lenient statistical thresh- olds (Figure 1). Activity selective for action verbs, relative to state verbs, was found posterior to verb-selective clusters. We then tested for differences between verbs and nouns that referred to states and events within functionally de- fined ROIs. Verb-selective LTC regions, defined by their preference for action verbs over object nouns, also re- sponded selectively to state and event verbs when con- trasted with state and event nouns (e.g., “he runs” vs. “the run”). Furthermore, neither MTG/STS nor pMTG re- sponded more to events than states. Instead, in the MTG/STS region, the opposite result was found, with stron- ger responses to states than events. Together, these results show that selectivity for action verbs in LTC cannot be fully accounted for by semantic differences between actions as agentive dynamic events and objects as concrete three- dimensional entities. Although verb selectivity in LTC may not directly reflect the retrieval of action concepts, as shown in this study, it is probably no coincidence that both verbs and concep- tual action knowledge recruit the left posterior MTG (Kemmerer et al., 2012; Campanella et al., 2010; Kalenine et al., 2010; Martin, 2007; Tranel et al., 2003; Martin et al., 1995). Considering the fact that most verbs describe ac- tions, it would be efficient for the brain to represent verbs close to regions representing action knowledge, as shown here (Figure 1). Importantly though, the present data sug- gest that verb processing and action knowledge, although represented nearby, are distinct processes. What could be the critical difference between verbs and nouns that drives activity in the left LTC? One possi- bility is that this region is involved in morphosyntactic processing needed for the correct inflectional form of verbs (Tyler et al., 2008; Tyler, Bright, Fletcher, & Stamatakis, 2004). Although we cannot exclude this possibility, it seems unlikely given that verb inflection was irrelevant to the pre- sent memory task. Further evidence against this possibility is provided by experiments involving semantic judgments of uninflected verbs and nouns that have found a verb effect in left LTC (Bedny et al., 2008, 2012). Alternatively or addi- tionally, LTC may be involved in the verb property of linking arguments within a sentence (Shetreet, Friedmann, & Hadar, 2010; den Ouden, Fix, Parrish, & Thompson, 2009; Thompson et al., 2007; Wu, Waller, & Chatterjee, 2007): Verbs specify that an agent does something (“he jumps”), has a certain property (“he stinks”), or does something with something else (“he builds houses”). Prototypical nouns lack this kind of predicative property (Lyons, 1977). A surprising finding was the increased activation to states relative to events in the left MTG/STS, considering that nearby regions (though typically in the right hemi- sphere) have been implicated in the processing of biolog- ical motion (Peelen, Wiggett, & Downing, 2006; Vaina, Solomon, Chowdhury, Sinha, & Belliveau, 2001; Allison, Puce, & McCarthy, 2000; Grossman et al., 2000). This dif- ference was particularly pronounced for verbs (Figure 2), with higher activation for state verbs like “he exists” than event verbs like “he chases.” Interestingly, a somewhat related preference for low-motion mental action verbs (e.g., to think) relative to high-motion action verbs (e.g., to kick) was observed in previous work (Bedny et al., 2008, 2012; Grossman et al., 2002). What could underlie this difference? One possibility is that the greater response for state verbs as compared with event verbs is related to the atypical semantic role of the subject: The subject of a state verb has the semantic role of undergoer, whereas typical subjects are agents (Van Valin & LaPolla, 1997; Givón, 1984). Another possibility may be that verbs that are distant from prototypical verbs (i.e., those that are less like agentive dynamic actions) recruit LTC more strongly. Reduced verb selectivity in LTC was found for the stimuli used in Experiment 1 as compared with those used in Experiment 2. One difference between the experiments was the use of animate subjects in Experiment 1 (“he” or “she”) and inanimate subjects (“this”) in Experiment 2. Given the significant verb selectivity observed for the local- izer conditions (action verbs vs. object nouns), which con- sistently used animate subjects, the relatively weak verb selectivity in Experiment 1 is unlikely to be related to the use of animate subjects. Perhaps more importantly, the nouns in Experiment 1 always had a verbal root, whereas those in Experiment 2 did not. Thus, nouns with verbal roots may partly activate verb-selective regions in LTC. Future studies should investigate this possibility more systematically. In contrast to several previous studies (e.g., Willms et al., 2011; Palti, Ben Shachar, Hendler, & Hadar, 2007; Shapiro et al., 2006), we did not observe significant verb-selective activation in left pFC at the corrected statistical threshold, although our sample size (N = 24) was larger than most of these previous studies that similarly contrasted action verbs with object nouns. This difference may relate to our task, which consisted of the simple memorizing of phrases and did not require task-relevant processing of morphological features as in some previous studies ( Willms et al., 2011; Peelen, Romagno, and Caramazza 2101 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f . t / u s e r o n 1 7 M a y 2 0 2 1 Palti et al., 2007; Shapiro et al., 2006). Indeed, two previous studies in which the task (a semantic relatedness task) similarly did not require morphosyntactic processing, verb-selective activity was found in the left pFC in only one of the studies (Bedny et al., 2008) whereas it was found in the left LTC in both studies (Bedny et al., 2008, 2012). In summary, this study showed that verb selectivity in LTC is not restricted to action verbs but is similarly present for nonaction (state) verbs contrasted with semantically matched nouns. In addition, the absence of a positive dif- ference between events and states suggests that verb selec- tivity in LTC is not related to the retrieval of event concepts. Although these results do not explain the critical com- ponent driving verb selectivity in LTC, they significantly re- duce the range of possibilities by excluding an explanation solely related to the retrieval of conceptual action knowl- edge. Equally important, given that the verb-selective effect in LTC cuts across semantic verb categories, it implies a general grammatical class effect that is not reducible to spe- cific semantic properties (such as action or event features). This leaves two possible organizational principles. One possibility is that the lexical distinction captured in LTC is a formal morphosyntactic property: a bundle of formal fea- tures that jointly serve to distinguish between nouns and verbs as morphosyntactically determined categories. The other possibility is that the distinction captured in LTC is that between words that typically do (verbs) and words that typically do not (nouns) have a predicative function. Predicates/verbs specify the types of roles or relations that arguments/nouns may take in a given situation, and as such can be thought of as a semantic rather than a syntactic category. This would be in line with results showing that LTC is implicated in conceptual processing. However, although we suspect that the latter possibility best captures the verb-selective activation in LTC, we still lack direct evi- dence for this claim, and we must await further theoretical analysis and experimental investigation before we can decide this issue conclusively. APPENDIX All the phrases used in the two experiments, with English translation. Also provided are the mean ratings for age of acquisition (higher ratings indicate older age of acquisi- tion), familiarity (higher ratings indicate higher familiarity), and imageability (higher ratings indicate higher imageabil- ity), separately for each condition. Standard deviation across participants is provided in brackets. Appendix. State verbs Experiment 1 Egli merita [he deserves] Lei preferisce [she prefers] Lei dissente [she disagrees] Egli crede [he believes] Experiment 2 Ciò include [this includes] Ciò implica [this implies] Ciò costa [this costs] Ciò riguarda [this concerns] Lei presuppone [she presumes] Ciò significa [this means] Lei eccelle [she excels] Egli vale [he is worth] Lei piace [she is liked] Egli risiede [he resides] Lei esiste [she exists] Lei eguaglia [she equates] Egli puzza [he stinks] Egli conosce [he knows] Egli teme [he fears] Lei manca [she lacks] Egli resta [he stays] Lei desidera [she desires] Egli giace [he lies (down)] Egli possiede [he owns] Lei sa [she knows] Ciò caratterizza [this characterizes] Ciò contiene [this contains] Ciò vale [this is worth] Ciò dista [this is far from] Ciò depende [this depends] Ciò piace [this is liked] Ciò esiste [this exists] Ciò manca [this lacks] Ciò puzza [this stinks] Ciò abbonda [this abounds] Ciò dispiace [this is not pleasant] Ciò sussiste [this subsists] Ciò comporta [this entails] Ciò incombe [this is incumbent] Ciò appartiene [this belongs] 2102 Journal of Cognitive Neuroscience Volume 24, Number 10 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j . f t / u s e r o n 1 7 M a y 2 0 2 1 Appendix. (continued ) Age of acquisition Familiarity Imageability State nouns Age of acquisition Familiarity Imageability Event verbs Experiment 1 Experiment 2 3.3 (0.6) 5.8 (1.1) 3.5 (1.2) Il merito [the merit] Lʼequivalenza [the equivalence] Il dispiacere [the displeasure] Il bisogno [the need] Il desiderio [the desire] Il possesso [the ownership] La mancanza [the lack] La conoscenza [the knowledge] La dipendenza [the dependence] Il timore [the fear] Il sapore [the taste] 4.2 (0.8) 6.1 (0.7) 3.9 (1.5) La temperatura [the temperature] Lʼegoismo [the egoism] Il prestigio [the prestige] Il carisma [the charisma] Il sonno [the sleep] Lʼidentità [the identity] Lʼaspetto [the look] Il genere [the kind/genre] Lʼaffetto [the affection] Lʼattitudine [the aptitude] Il nesso [the link] Lʼignoranza [the ignorance] La dimensione [the dimension] Lʼodio [the hate] La gioia [the joy] Il valore [the value] Il costo [the cost] Lʼeccellenza [the excellence] La necessità [the necessity] La qualità [the quality] La maniera [the manner] La pace [the peace] Lʼindole [the temperament] Il calore [the heat] La penuria [the scarcity] Lʼappartenenza [the membership] La prerogativa [the prerogative] La somiglianza [the resemblance] La particolarità [the peculiarity] 3.6 (0.7) 6.0 (1.0) 3.8 (1.2) 4.7 (0.8) 5.8 (0.9) 4.0 (1.4) Egli distrugge [he destroys] Egli insegue [he chases] Ciò arriva [this arrives] Ciò funziona [this works] Egli consuma [he consumes] Ciò aiuta [this helps] Lei scrive [she writes] Lei mangia [she eats] Lei crea [she creates] Egli osserva [he watches] Egli chiude [he closes] Egli uccide [he kills] Egli racconta [he narrates] Egli cammina [he walks] Lei piange [she cries] Lei arriva [she arrives] Ciò avviene [this happens] Ciò produce [this produces] Ciò succede [this occurs] Ciò prosegue [this goes on] Ciò gira [this spins/turns] Ciò accade [this happens] Ciò appare [this appears] Ciò torna [this comes back] Ciò bagna [this wets] Ciò cambia [this changes] Peelen, Romagno, and Caramazza 2103 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t . / f u s e r o n 1 7 M a y 2 0 2 1 Appendix. (continued ) Age of acquisition Familiarity Imageability Event nouns Experiment 1 Egli viaggia [he travels] Lei corre [she runs] Lei gioca [she plays] Lei parte [she leaves] Lei torna [she comes back] Lei parla [she talks] Egli lavora [he works] 2.2 (0.6) 6.2 (1.1) 5.9 (1.1) Experiment 2 Ciò causa [this causes] Ciò coinvolge [this involves] Ciò trasforma [this transforms] Ciò scorre [this flows] Ciò segue [this follows] Ciò colpisce [this strikes] Ciò finisce [this finishes] 3.3 (0.9) 6.4 (0.5) 4.7 (1.3) La distruzione [the destruction] Il festival [the festival] Lʼinseguimento [the chase] Lʼattacco [the attack] Il consumo [the consumption] Lʼuccisione [the killing] Il ritorno [the return] Il viaggio [the journey] Il pianto [the cry] Lʼuso [the use] La rissa [the fight] La vacanza [the holiday] Lʼinchiesta [the inquiry] La scampagnata [the jaunt] La cerimonia [the ceremony] La gita [the trip] Il disastro [the disaster] Lʼescursione [the excursion] La partenza [the departure] La conferenza [the lecture/conference] Lʼarrivo [the arrival] Lʼesplorazione [the exploration] La corsa [the run] Il moto [the motion] Il matrimonio [the wedding] Il convegno [the conference] La conversione [the conversion] Il tirocinio [the training] La lotta [the fight] Lʼesplosione [the explosion] La camminata [the walk] Il funerale [the funeral] Lʼincidente [the accident] Lʼimboscata [the ambush] Il trasferimento [the transfer] Il pellegrinaggio [the pilgrimage] Age of acquisition Familiarity Imageability Action verbs Lʼerosione [the erosion] Lʼazione [the action] 3.2 (0.6) 5.6 (1.1) 5.3 (1.0) Egli salta [he jumps] Lei colpisce [she hits] Lei combatte [she fights] Lei passeggia [she walks] Lei abbraccia [she hugs] Egli nuota [he swims] Il congresso [the convention] Lo spettacolo [the show] 4.8 (0.7) 5.9 (0.9) 6.0 (0.8) Same as in Experiment 1 2104 Journal of Cognitive Neuroscience Volume 24, Number 10 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t f . / u s e r o n 1 7 M a y 2 0 2 1 Appendix. (continued ) Age of acquisition Familiarity Imageability Object nouns Experiment 1 Lei disegna [she draws] Egli marcia [he marches] Egli ruba [he steals] Egli balla [he dances] Lei rompe [she breaks] Egli fugge [he runs away] Lei attraversa [she crosses] Egli canta [he sings] Lei costruisce [she builds] Lei trascina [she drags] Egli beve [he drinks] Egli legge [he reads] Lei applaude [she claps] Egli spinge [he pushes] 2.4 (0.5) 5.9 (1.5) 6.1 (1.2) La matita [the pencil] La pentola [the pan] La lampadina [the lightbulb] La poltrona [the armchair] La bacinella [the bowl] La bottiglia [the bottle] Il cuscino [the pillow] Il quaderno [the notebook] Il secchio [the bucket] Lo spillo [the pin] La maniglia [the handle] La terrazza [the terrace] Il biscotto [the biscuit] La tovaglia [the towel] Il cucchiaio [the spoon] Il campanile [the steeple] Il cassetto [the drawer] Il bicchiere [the glass] La camicia [the shirt] Lo sgabello [the stool] Experiment 2 D o w n l o a d e d f r o m Same as in Experiment 1 l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 4 / 1 2 0 4 / 2 1 0 0 9 / 6 2 1 0 9 9 4 6 4 / 5 1 1 9 7 7 o 8 c 2 n 7 _ 9 a / _ j 0 o 0 c 2 n 5 7 _ a p _ d 0 0 b 2 y 5 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t / . f u s e r o n 1 7 M a y 2 0 2 1 Age of acquisition Familiarity Imageability 2.1 (0.6) 6.1 (1.5) 6.6 (1.1) Peelen, Romagno, and Caramazza 2105 Acknowledgments We thank Silvia Ubaldi for help with data collection. 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