“Shall I compare thee”: The neural basis of literary awareness, and its benefits to

cognition

Noreen O'Sullivan, Philip Davis, Josie Billington, Victorina Gonzalez-Diaz, Rhiannon

Corcoran*

Department of Psychological Sciences, Institute of Psychology, Health, & Society, University

of Liverpool

*Corresponding author

rhiannon.corcoran@liverpool.ac.uk

+44 151 795 5365

B211, 2nd Floor Block B Waterhouse Buildings,

1-5 Brownlow Street, Liverpool, L69 3GL, UK

Running title: literary awareness

Word count: 6,715

Literary awareness

Abstract

FMRI was used to explore the neural and cognitive basis of literary awareness: the

capacity to process and derive meanings in complex text. A 2 x 2 design was used through

which 24 participants read short sections of poetic and prosaic text during scanning that

required significant reappraisal during reading, or did not require such reappraisal. Following

this, participants rated each piece on its ‘poeticness’ and the extent to which it prompted a

reappraisal of meaning during reading, providing subjective measures of poetic recognition

and the need to reappraise meaning. The substantial shared variance between these 2

subjective measures provided a proxy measure of literary awareness, which was found to

modulate activity in regions comprising the central executive and saliency networks. We

suggest that enhanced literary awareness is related to increased flexibility of internal models

of meaning, enhanced interoceptive awareness of change, and an enhanced capacity to reason

about events. In addition, we found that the residual variance in the measure of poetic

recognition modulated right dorsal caudate activity, which may be related to tolerance of

uncertainty. These findings are consistent with evidence that relates reading to improved

mental wellbeing.

Key words: poetry, caudate, default-mode network, saliency network, central executive

network

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Reading literature is encouraged as an activity in various health and community

settings as it is believed to be of therapeutic benefit. This is particularly so for shared-reading

(SR). In the UK, SR has been found to encourage solo reading and to facilitate

improvements in mental wellbeing (e.g. Dowrick, Billington, Robinson, et al., 2012;

Longden, Davis, Billington et al. 2015). However, very little is known about why reading

might benefit wellbeing. In this study, our starting premise is that increased reading is

associated with changes in cognition that promote wellbeing. For example, increased reading

of literature may increase one’s ability to engage the cognitive mechanisms that underpin the

representation of meaning. In individuals who struggle with mental health difficulties,

meaning derivation tends to be biased by dysfunctional and rigid beliefs. The encouragement

to consider alternative and nuanced meanings through active reading of texts may generalize

to encourage more adaptive, less rigid and biased meaning derivation in everyday life; thus

facilitating mental health and wellbeing. Here, we aimed to establish the neural basis of

mechanisms that support the capacity to consider, manipulate, and derive meaning in

complex text, a capacity that we refer to as literary awareness.

1.1 A cognitive perspective of meaning derivation, and its relationship to event complexity

The term ‘situation model’ describes the internally simulated contents of an

inference(s) attributable to a set of events (Kintsch, 1988; Kintsch & Rawson, 2005). The

contents of a situation model, which might include details related to time, space, causation,

motivation, and the protagonist (Zwann & Radvansky, 1998), are updatable in attempts to

generate a holistic, and accurate, interpretation of a set of events. Taking reading as an

activity that encourages the use of situation models (see Perfetti, Landi, & Oakhill, 2005, for

an overview of reading comprehension as a skill), the complexity of the text is likely to be

one factor that will determine the inherent nature of consequent models. In a simple text

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events may be wholly understood in relation to specific actions of the protagonist who is

motivated by easily recognizable circumstances in a singular point of time and space. In

contrast, in the complex texts offered by serious literature (fiction and poetry), the meaning

behind events can be multifaceted; the motivations of a protagonist may be unclear and

multiple, embedded in convoluted circumstances that enmesh past, present, and predicted

future.

The term ‘serious’ has a long history (The Oxford Professorship of Poetry was

established for “the advancement of more serious literature both sacred and human” [Henry

Birkhead]), and is widely used by writers and literary critics to distinguish literature that

addresses important human issues, such as those concerning social and moral behavior, from,

for example, ephemeral writings (comics, popular magazines) and genre fiction (e.g.

detective novels, ‘chick lit’; Dowrick et al., 2012). In order to appreciate meaning in serious,

or complex, texts, including poetry, the immersed reader needs to be alert to changes in the

direction of meaning, and focus their attention on maintaining a less rigid, more fluid

representation that can accommodate the overtures of a metaphor, and multiple meanings

woven in unusual turns of phrase. The characteristics of the text may be one underlying

factor that contributes to the efficacy of SR groups; individuals struggling with chronic pain

reported a preference for serious literature over simple texts, as these encouraged them to

think more deeply, and such reports coincided with a reduced experience of pain during

absorption in complex texts (Billington, Humphreys, Jones, et al., 2014).

We propose that in order to understand texts of increased complexity: 1) situation

models would need to be less rigid, and more flexible to accommodate varying related

meaning threads; 2) awareness of subtle differences in meanings would need to be increased;

and 3) social reasoning skills would need to be used to a greater extent to arbitrate between

the various possibilities. To summarize these necessary processing attunements we use the

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term ‘literary awareness’. In order to test our hypothesis that complex texts affect meaning

derivation in this way, we contrasted the meaning derivation of poetic (complex) and prosaic

(less complex) pieces. This hypothesis rests within a wider rubric that increased literary

awareness, with its accompanying changes to cognition, may form a substantial component of

the mechanistic basis of the efficacy of SR groups.

1.2 A neural perspective of meaning derivation, and its relationship to complexity

FMRI has been extensively used to explore meaning derivation in reading. Meta-

analyses across studies of text comprehension reveal a network of regions including nodes of

the default mode network (DMN) - the posterior cingulate cortex (PCC), dorsomedial

prefrontal cortex (dmPFC), and the anterior temporal lobe (ATL; Ferstl, Neumann, Bogler, &

von Cramon, 2008; Ferstl, Rinck, & von Cramon, 2005). While activation in PCC is thought

to reflect the extent to which a situation model has been updated, ATL is believed to store the

narrative of a situation model. The dmPFC forces attention to settle on a narrative for a

particular simulation that is heavily influenced by memory.

In addition to PCC and ATL, other regions that co-activate with the dmPFC during

language tasks include middle temporal gyrus (MTG), temporoparietal junction (TPJ) and

surrounding regions (BA39, 40), inferior frontal gyrus (IFG, BA 45/47), and ventromedial

prefrontal cortex (vmPFC, BA 11; Ferstl et al., 2008). Activation of TPJ and surrounding

areas in ventrolateral parietal is believed to indicate reflexive updating of situation models in

line with information retrieved from memory (Cabeza, Ciaramelli, Olson, & Moscovitch,

2008; Ciaramelli, Grady, & Moscovitch, 2008). Activation in inferior frontal gyrus (IFG) is

thought to maintain contextual separation between representations that are similar.

Consistent with this, IFG is active during the processing of metaphors (Eviatar & Just, 2006;

Lee & Dapretto, 2006; Rapp, Leube, Erb, et al., 2004; Shibata, Abe, Terao, & Miyamoto,

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2007; Stringaris, Medford, Giampietro, et al., 2007). Activation in ventromedial prefrontal

cortex likely reflects the motivational significance of the developing meaning (Cardinal,

Parkinson, Hall, & Everitt, 2002; Wallis, 2007).

While this extended semantic network is sufficient for simpler texts, more dynamic

and evolving texts would be likely to evoke the additional contribution of lateral anterior

prefrontal cortex (AP; BA 10). This area is thought to be involved in construing relationships

between less directly related words/meaning threads (Bunge, Wendelken, Badre, & Wagner,

2005; Green, Fugelsang, Kraemer et al., 2006; Green, Kraemer, Fugelsang, et al., 2010;

Monti, Osherson, Martinez, & Parsons, 2007; Monti, Parsons, & Osherson, 2009; Rutter,

Kröger, Stark, et al., 2012; Wendelken, Nakhabenko, Donohue, et al., 2008). Consistent with

a medial-to-lateral distinction in the role of AP in cognition, AP activation is increasingly

medial when decisions relate to encoded memories, and increasingly lateral when decisions

relate to novel representations (Baird, Smallwood, Gorgolewski, & Margulies, 2013). Areas

that become active as task difficulty increases, namely dorsolateral prefrontal (dlPFC) and

superior and posterior parietal regions that underlie the working memory network (Corbetta,

Patel, & Shulman, 2008), would also be expected to be necessary in supporting meaning

derivation from complex texts.

In relation to updating a model of meaning, we hypothesize roles for the dorsal

caudate, and the salience network (SN; Seeley, Menon, Schatzberg, et al., 2007). Activity in

the left dorsal caudate has been related to the deductive application of a rule from a set of

arguments (Melrose, Poulin, & Stern, 2007). Along with the putamen and lateral prefrontal

regions BA9 and 44, it is active across tasks that require categorical reasoning (Prado,

Chadha, & Booth, 2011). Activity in dorsal caudate during reasoning tasks may stem from

attunement to changes in an online representation. In simple reinforcement tasks, for

example, activity in the dorsal caudate increases when a behavioral choice becomes

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associated with a better outcome (Delgado, Locke, Stenger & Fiez, 2003; Knutson & Cooper,

2005; O’Doherty, Dayan, Schultz, et al., 2004; O’Sullivan, Szczepanowski, El-Deredy et al.,

2011). By extension, in reasoning tasks, caudate activity may reflect the better understanding

derived from the integration of an updated meaning into a situation model. Consistent with

this hypothesis, we previously found that activity in the left dorsal caudate was triggered by

Shakespearean functional shifts that required individuals to reason about a familiar word, and

its context, in a novel way (Keidel, Davis, Gonzalez-Diaz et al., 2013).

The SN is made up of regions connected to interoceptive awareness, including the

right fronto-insula, anterior insula, dorsal ACC/paracingulate cortex, and superior temporal

pole, but with extensions to amygdala, ventral striatopallidum, dorsomedial thalamus,

hypothalamus, pariaqueductal gray, and substantia nigra/ventral tegmental area (Seeley et al.,

2007). The network functions to direct cognitive resources towards sensory input, or online

representations, of motivational significance. In carrying out this function, the anterior insula

(AI), in particular, it is thought to play a role in switching from the DMN to the central

executive network (CEN) when motivation is entrained by an external event(s) (Menon &

Uddin, 2010; Uddin, Supekar, Ryali, & Menon, 2011). In the context of updating a model of

meaning, activation of the SN may coincide with activation in AP. Accepting a medial-to-

lateral distinction in AP function, coincident activation in AP would be expected to be more

lateral if the developing meaning model is influenced to a stronger extent by external content,

and more medial if the meaning model is influenced by memory (Baird et al. 2013; Bunge et

al. 2005; Green et al. 2006, 2010; Monti et al. 2007, 2009; Rutter et al. 2012; Wendelken et

al. 2008).

In the one previous published study that has directly contrasted poems with prose

(Zeman, Milton, Smith, & Rylance, 2013), regions related to the processing of poetry

included right posterior and mid-cingulate gyrus, left superior temporal gyrus (BA21), right

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temporal pole (BA 38) and bilateral hippocampi. Regions modulated by level of ‘literariness’

included left precentral gyrus, cingulate gyrus, left thalamus and caudate body, and left

cerebellum. Notably, the participants were experts and the selection of poems was familiar to

them, likely accounting for the absence of key regions hypothesized to be necessary in

building and updating new models of meaning. In contrast, when unfamiliar metaphoric lines

of poetry are contrasted with familiar sentences, left dlPFC and posterior middle temporal

gyri are recruited to a greater extent. This finding is consistent with the greater cognitive

challenges of poetry including, for example, greater control over the semantic processing of

sentences (Mashal, Faust, Hendler, & Jung-Beeman, 2009).

1.2 Current study

In the current study we scanned participants as they read and considered unfamiliar

pieces of poetry and prose of four lines in length. To enable us to systematically explore the

updating of models of meaning, the final line in half of the pieces was unexpected, triggering

a reappraisal of meaning. Neural responses to poetic and prosaic pieces were contrasted. At

a minimum, deriving meaning from poetry was expected to be more challenging than

deriving meaning from prose; thus we expected increased activity in IFG and dlPFC in

relation to processing poetic texts. Neural activity related to unexpected (where the final line

triggered a need to re-appraise) and expected (where the final line was consistent with the

meaning developed from lines 1-3) pieces was also contrasted. Here we anticipated enhanced

activity in AP and dorsal caudate in response to unexpected pieces.

Reading and comprehension are skills characterized by vast individual differences.

Indeed, we hypothesize that exposure to literature in the context of SR groups have their

effect on wellbeing by scaffolding individuals’ ability to derive meaning from complex sets

of circumstances—building literary awareness. In order to assess the extent to which

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individual differences modulated the representation of meaning, we computed a number of

behavioral regressions to include in relevant fMRI contrasts. It was assumed that individuals

who are more aware of the nuances in complex text would be better at discriminating

between simple and complex texts, and furthermore, the extent to which they can do this

would modulate their capacity to derive meaning in text generally. Following the scanning

session, participants rated all of the stimuli on the extent to which they perceived them as

poetic. From this we derived a measure of ‘poetic recognition’ by subtracting poetic ratings

for prosaic pieces (simple texts) from poetic ratings for poetry pieces (complex text).

A substantial amount of the variance in individuals’ capacity to discriminate between

complex (poems) and simple (prose) texts would be expected to relate to an underlying

ability to notice changes in meaning, and represent alternative meanings between related

meaning threads. We therefore also asked participants to rate the extent to which they

perceived a need to reappraise the developing meaning on reading the final line. From this, a

measure of ‘need for reappraisal’ was calculated by subtracting the perceived need to

reappraise meaning from pieces that ended as expected (expected pieces) from those that

ended unexpectedly (unexpected pieces). The variance shared between ‘poetic recognition’,

and ‘need for reappraisal’, was computed to act as a summary, proxy measure of literary

awareness. We chose the approach of taking the shared variance between poetic recognition

and need for reappraisal as our measure of literary awareness because the simple measure of

poetic recognition was likely to be influenced by other, unmeasured sources of individual

difference that would be used in the identification of verse (e.g. attunement to the musicality

of poetic text). If, as we expect, our measure of literary awareness reflects the skills that

support the flexible and iterative updating of meaning, tied to the timely engagement of the

SN, then we would expect the measure of literary awareness to modulate activity in dorsal

caudate, AP (medial and lateral), and cingulate, insula, and thalamic nuclei specifically.

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2. Method

The University of Liverpool Research Ethics Committee approved the study, and

participants’ consent was obtained according to the Declaration of Helsinki.

2.1 Participants: To ensure that participants were able to consider and reflect on the

brief, but complex, verbal stimuli we were using under timed conditions in the scanner, we

selected a group of individuals who we knew would be familiar enough with the

contemplation of poetry to enable them to meaningfully process the pieces. At the same time

we did not want to sample literature scholars where processing differences associated with

‘expertise’ may have been found. With this in mind, we recruited 24 right-handed first or

second year undergraduate English students (16 female; age-range 18-30) to take part in this

study.

2.2 Stimuli: Poetic pieces were chosen by the literature scholars in the research team

according to the needs stipulated by the agreed research questions. These were all taken from

published, though not widely familiar, poems. The prosaic pieces were constructed by the

research team based upon the theme and meaning embedded in poetic pieces that were not

included in the final selection of stimuli for scanning. All pieces were 4 lines long with

punctuation controlled across poetic and prosaic pieces. Poetic and prosaic pieces were

further subdivided into those that gave rise to a global meaning that could not be anticipated

based on the first 3 lines of the text (unexpected meanings requiring reappraisal of the piece),

and those with a global meaning that developed linearly during reading (expected meaning).

92 stimuli (46 poetic, 46 prosaic) were rated by a sample of 30 individuals (age-range, 16-65;

mix of educational backgrounds, with all individuals having had formal education to at least

age 16). Using 7 point scales, each stimulus was rated on confidence that the piece’s

meaning had been understood (unconfident-confident); the feeling that the piece generated

(negative-mixed-positive); the extent to which a piece had a different global meaning to that

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derived in the initial 3 lines (predicted-unpredicted meaning); and how poetic the piece was

felt to be (prose-poem).

24 poetic and 24 prosaic stimuli (12 expected and 12 unexpected for each; see Table

S2) were selected for use in the scanner. Each poetic piece was matched to a prosaic piece

both in valence and global theme. The number of words per piece did not significantly differ

between conditions (see Table S1). Stimuli of either a positive, negative, and ambiguous

emotional valence were equally represented in each condition (4 in each condition, in order to

control for the global emotional tone of the pieces read per scan). Examples of positive

themes included love and luck; examples of ambivalent themes included loss-related growth

and loneliness -related hope/faith; examples of negative themes included loss-related sadness

and danger. The same pieces were presented together in a scan for each participant, and the

order of blocks was counter-balanced across participants. Participants saw the stimuli during

the scans, and then again after scanning when they rated each piece.

2.3 Design and procedure: In the scanner, participants were asked to read and

consider each poetic and prosaic piece as best they could (reading), and to then reflect on the

meaning they had derived from the piece (reflection)—see Figure 1. 8 poetic and 8 prosaic

pieces were presented in random order in each of 3 functional scans. In each scan, four of

each linguistic category required reappraisal. To separate the signal related to the reading

and the reflection phases, and to prevent reflection on the piece prior to the reflection phase,

simple math equations were presented for 6 seconds before the onset of each piece. Each

functional scan lasted 14 minutes.

Following scanning, participants rated each stimulus on: 1) its perceived poeticness;

2) the sense of a need to reappraise the meaning of the piece after reading the final line; 3) the

extent to which they felt they understood its meaning; 4) its valence. ‘Poetic recognition’

was calculated by subtracting poetic ratings for prosaic pieces from poetic ratings for poetry

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pieces. ‘Need for reappraisal’ was calculated by subtracting the perceived need to reappraise

meaning from pieces that ended with an expected meaning (expected pieces) from those that

ended unexpectedly (unexpected pieces). Higher ratings on a 1-7 likert scale were indicative

of increased poeticness, and increased need to reappraise meaning.

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Figure 1 about here

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2.4 Data acquisition: Functional images were acquired using a 3 T Siemens Trio MRI

scanner with a gradient echoplanar sequence (TR = 2000 ms, TE = 30, flip angle = 90°).

Each volume consisted of 33 tilted axial slices co-planar with AC-PC (3 mm thick, 0.3 mm

slice gap; FOV 210 x 210; Matrix size 64 x 64) that provided near whole brain coverage. 432

volumes were collected per scan, with slices being acquired sequentially. Anatomical T1-

weighted images were acquired with the following parameters: TR = 500 ms, TE = 10, slice

thickness = 4 mm at a 90° flip angle. Head motion was limited using padding.

2.5 Data analysis: Data were analyzed using the FSL package, version 5.0.1.

Preprocessing steps applied to each functional scan included slice-timing correction to the

first slice, extraction of noise components in the data via Melodic ICA (using the criteria of

Kelly, Alexopoulos, Wang, et al., 2010, as a guide in selecting components), high pass

filtering that was estimated in individual scans, and spatially smoothing with a FWHM of

5mm. Individual functional scans were aligned to their anatomical counterpart using a linear

transformation method optimized with the Boundary-Based-Registration method, and were

subsequently standardized to the 2 mm MNI152 brain using a linear transformation method

with 12 degrees of freedom.

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Functional data in each scan were fit with an event-related general linear model that

was estimated through generalized least squares (i.e. OLS after whitening). Time stamps

were convolved with the standard canonical gamma function implemented in FSL (lag, 6 s;

width, 3s). Two models were fit to the linguistic stimuli. The first included 8 events from

the combination of task phase (read and reflection), linguistic category (poetic and prosaic),

and need for reappraisal (expected and unexpected). The time stamps corresponded to the

onset of each piece during reading and reflection. Four contrasts were of interest in relation

to the hypotheses: the contrast between poetic and prosaic stimuli during reading, and during

reflection (see supplementary material for conjunction contrasts).

The second model included 12 events. The 4 additional events in this model

corresponded with the onset timestamps of the 4th sentences in each condition (linguistic

category x reappraisal; separate events were included for the first 3 sentences of each

condition). Two contrasts were of interest in relation to the hypotheses: the contrast between

the 4th lines in pieces that required reappraisal (unexpected) relative to those that did not

(expected). Temporal derivatives were included in both models, and temporal filtering was

applied. Voxels were uncorrected for multiple comparisons at a threshold of p < .05 in each

individual scan.

Contrasts were averaged across runs within participant, with the subsequent averages

being entered in mixed-effect group analyses in which the parameters in all voxels close to

threshold, as established following a fast approximation of the model, were estimated using

Metropolis-Hastings Markov Chain Monte Carlo sampling to lead to the best estimates of

both fixed and random effects. Contrasts were cluster corrected at a z threshold of 2.3, and a

p-value of 0.05. Group contrasts were subsequently re-run to include individual differences

variables as regressors. The contrasts that were relevant in relation to the hypotheses

included modulation of each task phase (read and reflection), independently of linguistic

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category, modulation of contrasts in which prosaic activity was subtracted from poetic

activity, and modulation of the contrast related to reappraisal (specific to 4th lines; expected

versus unexpected).

3. Results

As expected, poetic pieces were rated as more poetic than prosaic pieces (poetic M =

5.56, SD = .52; prosaic M = 3.30, SD = .81; t (23) = 12.56, p < .001), and participants were

less confident in having understood the meaning of poetic pieces, (poetic M = 4.99, SD =

1.07; prosaic M =5.92, SD = 1.14; t (23) = 6.78, p < .001). Participants were generally

accurate at discriminating between pieces requiring a reappraisal of meaning from those that

did not, showing that participants were consciously aware of the need for reappraisal

(reappraisal M = 4.18, SD = .45; no reappraisal M = 3.62, SD = .42; t (23) = 4.75, p < .001).

There was no difference in participants’ reappraisal ratings for poetic and prosaic pieces

(poetic M = 4.00, SD = .41; prosaic M = 3.80, SD = .49). There was significant shared

variance between participants’ poetic recognition and need for reappraisal scores—r (24)

= .617, p = .001, with the predicted variance in poetic recognition from reappraisal (the

shared variance) providing the proxy measure of literary awareness. Residual scores in

poetic recognition (controlling for reappraisal), and residual scores in reappraisal (controlling

for poetic recognition) were also calculated for inclusion in the subsequent fMRI contrasts.

Although participants were less confident in having understood the meaning of poetic

relative to prosaic pieces, confidence was unrelated to literary awareness or need for

reappraisal and poetic recognition (ps > .38). This is important because it demonstrates that

perceived difficulty is not determining variability in these measures. Table 1 presents the

findings related to the 5 key contrasts.

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3.1 Contrasting poetic pieces with prosaic pieces

3.1.1 Reading: During the reading phase, poetic pieces differentially recruited more

lateral frontal and temporal/occipital regions than prosaic pieces. These areas included

bilateral precentral and IFG, and right-sided dlPFC. Activation was more extensive on the

left, extending into the anterior insula, and beyond to the temporal pole. In contrast, a cluster

in dmPFC showed reduced activation during reading of poetic pieces, compared to the

reading of prosaic pieces.

3.1.2 Reflection: During the reflection phase, differential activation in left IFG and

left LOC remained. The reduced activation associated with the reading of poetry seen in

dmPFC extended during reflection to include other areas loosely corresponding with the

DMN (Figures S2 & 3).

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Table 1 about here

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3.2 Contrasting the reading of unexpected to expected: the need for reappraisal

During reappraisal (prompted by the reading of unexpected final lines), clusters that

overlapped with reading in general were evident in medial frontal, posterior MTL, and in TPJ

(30%, 65%, and 88% respectively). These findings are consistent with a linear increase in

required activity during the reading of the unexpected final line. By contrast, and consistent

with a distinct role related to the need to reappraise, clusters that did not overlap with reading

in general appeared in left lateral SFG (BA9) and dorsal caudate, extending to anterior

thalamic nuclei (Figure S4), in response to the reading of the unexpected final line.

3.3 Modulation of activity by poetic recognition, reappraisal, and literary awareness

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Table 2 about here

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Table 2 shows how activity in response to reading and reflecting in general, followed

by the reading of and reflecting upon poetic stimuli specifically, was modulated by the

individual difference variables. During both the reading and the reflection phases of both

poetic and prosaic pieces, the proxy measure of literary awareness modulated a similar,

large, number of regions—see Figure 2. Lateral frontal areas included AP, dlPFC, IFG, and

precentral gyri, with activation being distinctly more right-sided in dlPFC and IFG.

Activation spanning regions within temporal/occipital boundaries was also stronger on the

right, in addition to activation in SMG. Similar levels of bilateral activity were observed in

the dACC, the insula, the basal ganglia, and the thalamus (extending to the brain stem). The

cerebellum was the only region in which bilateral activity was more left-sided. These

modulated clusters did not overlap with those active during reading and reflecting in general.

Note, we had not specifically hypothesized hemispheric differences in terms of modulated

region engagement; therefore the comparisons across hemispheres in Table 2 are reported in

order to facilitate better predictions in future work, rather than being an authoritative outcome

in their own right.

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Figure 2 about here

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3.3.1 Specific to Reading Poetry: During reading of poetic pieces specifically,

residual variance in poetic recognition (poetic recognition > subjective need to reappraise)

modulated a single cluster that spanned from right dorsal caudate to dACC, and further to

medial and lateral AP, see Figure 3.1. Right caudate activity in this cluster mirrored the

caudate activity of the left-sided cluster related to reappraisal (unexpected > expected)—see

Figure 3.3. About half of the left caudate voxels active during reappraisal (unexpected >

expected) were modulated by literary awareness and reappraisal during reading (57%). As

Figure 3.2 shows, when reappraisal was most necessary (unexpected > expected), meaning

derivation became increasingly dependent on left rostral caudate.

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Figure 3 about here

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3.3.2 Specific to Reflecting on Poetry: During reflection of poetic pieces specifically,

literary awareness modulated a number of clusters—see Figure 4. Characteristic areas of the

DMN included PCC, precuneus, and dmPFC. Lateral frontal areas included AP, with

stronger activation on the left, IFG stronger on the right, and bilateral orbitofrontal cortex

(OFC) extending to the insula. ACC was also active. Temporal areas that were left-sided

included those overlapping temporal/occipital boundaries, posterior MTG, LOC and FG.

Activation extended to the hippocampus, parahippocampal gyrus, and putamen. In parietal

cortex, bilateral activity in SMG was evident. Finally, there was a left-sided cluster in the

cerebellum.

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Figure 4 about here

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4. Discussion

Our aim was to explore the neural basis of literary awareness—the capacity to derive

meaning in complex texts such as poetry. We were interested in this because we hypothesize

that exposure to literary texts can support a more fluid and flexible representation of

meaning, in order to allow for: 1) multiple potential truths to be weighted with similar levels

of curiosity, 2) changes in meaning to be noticed more easily, and 3) online meanings to be

more easily updated as additional information becomes available. We draw a parallel

between the non-linear process a reader goes through in reading a complex text, and the mix

of uncertainties, choices, blunderings, successes, and insights that we all live through on a

daily basis. This comparison extends to the premise that flexing the mechanisms that support

meaning derivation in the context of active reading may entrain individuals towards more

responsive conceptualizations of meaning in their own lives. Our findings support this

premise in that they demonstrate how our proxy measure of literary awareness modulated

regions associated with the ability to represent and manipulate fluid, flexible meanings.

4.1 The neural basis of our measure of literary awareness

The co-activation of dlPFC (BA 46), IFG, temporo-occipital regions, and ATL during

reading of poetry relative to prose is consistent with poetic text requiring the representation of

multiple meaning threads (IFG), needing more focused attention during processing (dlPFC;

temporo-occipital regions) in order to generate a holistic model of meaning (ATL).

Deactivation in dmPFC, in the context of increased activation in dlPFC and AI, suggests that

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processing of poetic content requires a switch away from stored representations to build

meaning from a novel external source. Deactivation of multiple regions that typically co-

activate in the DMN during the reflection phase potentially points to the longer time needed

to establish a stable representation of meaning for poetic pieces. Consistent with this, a

continued increase in the extent of activation in IFG and LOC while reflecting on poetry is

indicative of participants appraising varying meanings.

Our proxy measure of literary awareness modulated a number of regions making up

the CEN (dlPFC, lateral parietal, IFG, AP, and dorsal caudate) and the SN (insula, ACC,

striatopallidum, thalamus, and brain stem). Co-activation of both of these networks suggests

that increased literary awareness is indeed related to an improved capacity to reason about

events in a responsive manner, and an increased interoceptive attunement to changes in

meaning. Additional activity in OFC is also consistent with situation models having an

external focus, this time in relation to the motivational significance of meaning, as opposed to

an internal focus, where motivational significance is derived through memory (Bouret &

Richmond, 2010). Finally, activation extended to ventral striatum/nucleus accumbens, a

region associated with the perception of rewarding information (O’Doherty et al., 2004);

thus, enhanced literary awareness may increase the extent to which contemplating meaning

is perceived to be rewarding. Notably this same set of regions was modulated during reading

of, and reflection on, both simple and complex texts. This suggests that attempts to improve

literary awareness through, for example, SR groups, should have the power to alter how

individuals derive meaning in life events generally, regardless of their complexity. It does

seem possible that the efficacy of SR groups in improving wellbeing could be moderated by

the extent to which this set of regions become engaged in deriving meaning.

Modulated activity in left dorsal caudate overlapped with caudate activity that was

evoked by unexpected meanings (unexpected > expected). Activation became increasingly

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rostral as the need to update meaning became more obvious. The augmented capacity to

update internal representations associated with increased literary awareness is consistent with

more flexible/responsive meaning models. Strikingly, residual variance in poetic recognition

(variance set apart from that which could be attributed to reappraisal) modulated right dorsal

caudate. Dissociable roles for the left and right dorsal caudate in relation to the updating of

representations have yet to be delineated. We surmise that whereas left caudate activity

reflects responsive updating of online representations, modulated activity in right caudate

may reflect individual variability in the overall tolerance of meaning uncertainty.

While reflecting on poetry in particular (more than simple texts), literary awareness

modulated concurrent activity in nodes of the DMN (PCC, precuneus, dmPFC), AP and IFG,

bilateral posterior parietal, and regions of the SN (insula, ACC, putamen). This suggests that

increased literary awareness is related to an increased ability to maintain a developing

meaning model (DMN) from complex material, supported by the ability to maintain

dissociable meaning threads (IFG) that are actively reasoned about (AP and bilateral parietal)

with a high level of awareness (SN). Although there remains work to be done on the extent

to which text complexity and seriousness might moderate the benefits of SR groups to

wellbeing, it has always been the intuition of literary experts involved that complex texts

from serious literature, would be of greater benefit than linear plot-based novels. In complex

texts, the meaning behind events travels through subtle shifts, sometimes leading to an

internal state where multiple possibilities are compatible. It is in this complexity that serious

literature has a greater capacity to emulate what it is really like to be human: the thoughts,

feelings, and behaviors of imperfect characters become understood in relation to their

sociocultural context. Increasing individuals’ ability to deal with human social complexity in

text may be the driver that enables more fluid, and better reasoned, situation models of their

own complex human issues.

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Literary awareness

Parietal activity was more posterior and dorsal to TPJ, the latter being recognized to

play a core role in reflexive access to memory (Cabeza et al., 2008; Ciaramelli et al., 2008).

It is possible that the spatial change in parietal activity reflects the effort to maintain internal

working models of meaning constructed from an external stimulus. However, concurrent

modulation of the precuneus is consistent with episodic memory retrieval and in particular,

the use of self-referential knowledge in supporting the production of inferences (see Cavanna

& Trimble, 2006, for a review). Activation of the area has also been reported in association

with efforts to build mental images to support holistic understanding during reading of poetry

(Mashal, Vishne, & Laor, 2014). Thus there is scope to consider that self-referential

knowledge of the world is used to augment access to meaning in text. Finally, modulation of

the hippocampus by literary awareness suggests that the nuances of meaning derived from the

text were consolidated to memory making them available for meaning searches in the future.

In relation to the efficacy of SR groups in improving wellbeing, the hippocampal activity

reflecting memory consolidation points to a potential pathway through which social

reasoning practiced in SR groups could come to support social reasoning in everyday life.

We considered it possible that the difficulty of deriving meaning in poetry, and not

our measure of literary awareness, might be the key explanatory factor relating to enhanced

or differentiated brain activity. However, individual differences in confidence to do with

‘getting the meaning’ did not systematically modulate brain activity. It seems, therefore, that

perceived difficulty is separable from the mechanisms that contribute to literary awareness.

4.2 From literary awareness to mental wellbeing

One of the key motivations for conducting this study was to explore how functional

brain imaging might help to elucidate the factors that underpin the ‘wellbeing’ effect of active

reading. We argue that at its core, poetic processing shares the same fundamental

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Literary awareness

mechanisms that underpin the processing of, and decision-making within, the fluid, non-

linear and ambiguous social world (e.g. Corcoran, 2001). As has been argued that the

processing of metaphors requires the same complex skills that underpin theory of mind

(Happe, 1995; Bottini, Corcoran, Sterzi et al., 1994). Consistent with this, increased reading

of literary fiction, and not popular fiction or non-fiction, has been linked to improved use of

both cognitive and affective aspects of theory of mind (Kidd & Castano, 2013). Hence, it can

be argued that the facility to iteratively appraise complex literary material mimics the

processes of effective dynamic social perception and cognition.

Contemporary neuroscientific theories represent our brains as highly efficient, but

fallible, prediction machines (Tenenbaum, Griffiths and Kemp, 2006). Expectation biases

derived through experience influence how information is processed moment-by-moment.

The prediction error signal exists to fine tune our information processing so that the meaning

that is ‘bound’ to a stimulus or an event can be updated to account for deviation from

expectancies (O’Doherty et al., 2004; Pessiglione, Seymour, Flandin, Dolan & Frith 2006).

At times, however, this usually reliable fast track to information processing leads to what can

appear to be irrationality (Tversky & Kahneman, 1974; Kahneman, 2012). Socio-cognitive

research into mental health difficulties suggests that both information processing biases and

adverse experiences ‘color’ our fast track processing, thus our view of the anticipated social

world, leading to distortions in predictive reasoning that, to others, appear as irrational

(Bentall, Corcoran, Howard, Blackwood & Kinderman, 2001; Corcoran & Frith, 2003;

Dannlowski, Stuhrmann, Beutelmann et al. 2012).

When very high value is attributed to a belief or action, prediction errors of minimal

size would not be expected to readjust the value of the belief or action to any meaningful

extent (Gradin, Kumar, Waiter et al., 2011; Moran, Owen, Crookes et al. 2008). In addition,

even when PEs are consciously registered, motivated reasoning that negates the PE as an

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Literary awareness

anomaly further serves to protect the high value of formerly reliable, tried and tested,

interpretations, attributions, beliefs, and actions (Kunda, 1990; Westen, Blagov, Harenski,

Kils & Hamann, 2006). When the value of maladaptive beliefs that sustain mental health

difficulties are protected in this way, decision-making and reasoning becomes ‘sticky’, rigid,

inflexible, and maladaptive, sustaining mental health difficulties (Corcoran, Cummins,

Rowse et al., 2006; Kahneman, Slovic, & Tversky 1982).

Part of the wellbeing efficacy of SR may come from pulling individuals away from

processing their struggles through tired and ineffective lenses and towards more diverse,

novel and effective reasoning options. Consistent with this, depressive beliefs have been

linked to dysfunctional connectivity in DMN, SN, and CEN (Hamilton, 2013; Menon, 2011),

in addition to reduced caudate volume and reduced dorsal- and ventral striatal activity

(Pizzagalli, Holmes, Dillon et al., 2009). Furthermore, increased literary awareness might

reduce ruminative thinking, readjusting connectivity between the DMN, SN, and CEN.

Consistent with this, lateral prefrontal activity is associated with the ability to avoid

responding on the basis of a belief, in favour of a logical response (De Neys, Vartanian, &

Goel, 2008; Goel & Dolan, 2003; Stollstorff, Vartanian, & Goel, 2012; Tsujii, Masuda,

Akiyama, & Watanabe, 2010; Tsujii & Watanabe, 2010). Increased literary awareness might

increase the tendency to search for an alternative meaning framework within which to situate

events, thus providing a scaffold to help in the replacement of dysfunctional with functional

beliefs that cannot so easily sustain worries, troubles and negative automatic thoughts.

In short, we contend that our results demonstrate that increasing literary awareness is

related to brain activity that supports greater capacity to process dynamic models of meaning.

If the reading of literature can encourage and train a more dynamic, fluid, and less rigid

reasoning style, then this will have intrinsic therapeutic benefits that should include

improved, adaptive social functioning.

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4.3 Limitations and future research

Our measure of literary awareness was largely influenced by one genre of complex

text: poetry. Although we sought to strengthen the measure by demonstrating the extent to

which variance in poetic recognition overlapped with the need for reappraisal, further work

could focus on whether our findings are limited to the representation of meaning in poetic

forms. In principle, however, we would expect that sensitivity to meaning complexity in any

text would modulate the same underlying mechanisms revealed here.

We chose a sample of English students in order to ensure that, in this first study,

baseline levels of literary awareness were high enough to guarantee some ability to recognize

and process poetry within the demanding confines of an fMRI activation study. While our

students were hardly literary scholars, the extent to which our findings reflect the relative

skills of our sample is unclear, and would need to be explored in future studies. The likely

influence of the backgrounds of our sample is reduced by the fact that all of the stimuli we

used in this experiment were unfamiliar to the participants.

Regardless of how participants were selected, it is likely that literary awareness is

related to level of general intelligence. In particular, we would expect this to be a type of

social intelligence, going beyond that typically measured in cognitive assessments of

intelligence and including aspects of social cognition such as perspective taking. Future

research of this nature should consider including measures of IQ and social intelligence. Our

sample had twice as many females as males, and we acknowledge that gender may play some

role in relation to the neural representation of literary awareness.

Using functional imaging in addressing hypotheses related to complex human abilities

increases the number of regions supporting task performance; thus increasing the likelihood

of a mismatch between the function that is attributed to particular region, and its true

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Literary awareness

function. In facing this issue, we created a well controlled 2 x 2 design, and supplemented

our analyses with individual difference variables. That said, with this caveat in mind, we are

hopeful that others will see the utility in assessing the extent to which measures of literary

awareness modulate functional activity during other tasks, both linguistic and non-linguistic.

Our brains are capable of complex abilities, and sometimes when we take a look at that

complexity, we see smaller meaning threads that, without the presence of a whole, would not

be seen; for example, the potential for trait and state (alternatively, tonic and phasic) aspects

to right- and left dorsal caudate function. Finally, we had no clearly justifiable reasons to

predict hemispheric differences modulated by literary awareness, another issue requiring

follow-up.

Despite these limitations, we argue that our findings suggest the manner in which

active reading can benefit wellbeing by encouraging a more responsive and flexible

information processing style that crosses the boundary between reading text, and reasoning in

every-day life. The experienced reader is aware of multiple meaning threads, and is vigilant

to the need to update meaning while building an iterative, yet holistic model of a poem or

plot; we are each readers and reflectors of our own and others’ developing plots: were we

better readers, would we not be better writers-?

5. Funding

This work was supported by in-house funding provided to the authors by the Institute

of Psychology, Health, and Society, University of Liverpool. The funding source had no

involvement in the study design, data collection and analysis, and the interpretation of the

data.

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Figure captions

Figure 1

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Literary awareness

Time-line for a single trial

Figure 2

T-map of areas modulated by literary awareness during reflection on both poetic and prosaic

stimuli (cluster corrected threshold of z = 2.3, p < .05 –increasing the threshold did not

substantially alter the pattern), spanning medial-to-lateral in the right hemisphere. Regions

included ACC, basal ganglia, thalamus, precuneus, SMG, temporal/occipital, insula, and

lateral frontal

Figure 3

T-map of areas modulated by residual poetic recognition during reading of poetic stimuli, far

left (1), including right caudate, extending to medial and lateral anterior prefrontal. Middle

(2) = caudate activity during reappraisal, specifically in red, compared with caudate activity

modulated by literary awareness during reading of both poetic and prosaic stimuli. Right (3)

= caudate activity during reappraisal, in red, compared to caudate activity modulated by

residual poetic recognition during reading of poetic stimuli

Figure 4

T-map of areas modulated by literary awareness during reflection on poetic stimuli (cluster

corrected threshold of z = 2.3, p < .05), spanning lateral-to-medial in the left hemisphere,

regions including medial temporal, supramarginal gyrus, lateral frontal, insula, putamen.

Right upper = right inferior frontal gyrus and supramarginal gyrus; right lower = medial

frontal and posterior cingulate

Figure S1

T-map of areas active during reading (cluster corrected threshold of z = 2.3, p < .05)

Figure S2

T-map of areas differentially active during reading of poetry (cluster corrected threshold of z

= 2.3, p < .05). Axial slices show bilateral temporal/occipital clusters, in addition to anterior

36

Literary awareness

temporal and insula activity; sagittal slices additionally show right-sided dlPFC,

IFG/precentral activity, and dmPFC deactivation

Figure S3

T-map of areas differentially active while reflecting on poetry (cluster corrected threshold of

z = 2.3, p < .05). Increased activation was observed in left IFG and LOC; deactivation was

observed in dmPFC, precuneous/posterior cingulate, and right-sided V1, lateral parietal, and

central opercular cortex

Figure S4

T-map of areas differentially active during reappraisal (cluster corrected threshold of z = 2.3,

p < .05). Regions included superior frontal gyrus, dACC, caudate, extending to thalamus,

and TPJ—all left-sided

Table captions

Table 1

37

Literary awareness

Statistics on brain regions that differentially responded to poetry during reading and

reflection, in addition to regions that differentially responded to reappraisal (unexpected >

expected). All clusters were corrected at z = 2.3, p < .05. The percentage of active voxels

that the reading contrasts shared with reading in general (poetry + prosaic), and that the

reflection contrasts shared with reflection in general are also shown, in addition to the

percentage of voxels that were modulated by literary awareness (LA). + = 57% overlap in the

caudate specifically; * = 35% overlap in precuneous specifically (PreCG = precentral gyrus;

IFGpo = inferior frontal gyrus pars orbitalis; FOC = frontal opercular cortex; a.ins = anterior

insula; TP = temporal pole; pMTG = posterior middle temporal gyrus; LOC = lateral

occipital cortex; FGpt & FGo = fusiform gyrus posterior temporal and occipital; pITG =

posterior inferior temporal gyrus; dACC = dorsal anterior cingulate; mAP = medial anterior

prefrontal; pSMG = posterior supramarginal gyrys; AG = angular gyrus; PCC = posterior

cingulate)

Table 2

Statistics on brain regions that were modulated by literary awareness (LA) and residual

variance in poetic recognition (PR). All clusters were corrected at z = 2.3, p < .05 (FOC =

frontal opercular cortex; ITG = inferior temporal gyrus; FG = fusiform gyrus; LG = lingual

gyrus; LOC = lateral occipital cortex; lm AP = lateral and medial anterior prefrontal; MFG =

middle frontal gyrus; OFC = orbitofrontal cortex; dACC = dorsal anterior cingulate; ins =

insula; hippoc = hippocampus; parahippoc = parahippocampal gyrus). Clusters in reading

and reflection conjunctions were extensive; therefore the volume of voxels for each of the

key regions within clusters was derived as a function of being left- or right-sided (prop. right

= proportion of voxels that were right-sided)

Table S1

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Literary awareness

Descriptive statistics on the number of words in each condition, and the average number of

words per line

Table S2

Stimuli used in the study

Table S3

Statistics on brain regions that responded similarly to poetic and prosaic stimuli during

reading and reflection. All clusters were corrected at z = 2.3, p < .05. The percentage of

active voxels that were modulated by the variance shared between literary awareness is

shown in the far right column (SFG = superior frontal gyrus; mAP = medial anterior

prefrontal; SubC = subcallosal cortex; STG = superior temporal gyrus; pSMG = posterior

supramarginal gyrus; AG = angular gyrus; TP = temporal pole; OFC = orbitofrontal cortex;

IFGpt = inferior frontal gyrus pars triangularis; Hippoc = hippocampus; Amyg = amygdala;

LG = lingual gyrus; MTG = middle temporal gyrus; IFGpt = inferior frontal gyrus pars

orbitalis; PCC = posterior cingulate

39