Running Head: Training interoceptive cortex Mindfulness meditation training alters cortical representations of interoceptive attention

One component of mindfulness training (MT) is the development of interoceptive attention (IA) to visceral bodily sensations, facilitated through daily practices such as breath monitoring. Using functional magnetic resonance imaging (fMRI), we examined experiencedependent functional plasticity in accessing interoceptive representations by comparing graduates of a Mindfulness-Based Stress Reduction course to a waitlisted control group. IA to respiratory sensations was contrasted against two visual tasks, controlling for attentional requirements nonspecific to IA such as maintaining sensation and suppressing distraction. In anatomically-partitioned analyses of insula activity, MT predicted greater IA-related activity in anterior dysgranular insula regions, consistent with greater integration of interoceptive sensation with external context. MT also predicted decreased recruitment of the dorsomedial prefrontal cortex (DMPFC) during IA, and altered functional connectivity between the DMPFC and the posterior insula, putative primary interoceptive cortex. Furthermore, meditation practice compliance predicted greater posterior insula and reduced visual pathway recruitment during IA. These findings suggest that interoceptive training modulates task-specific cortical recruitment, analogous to training-related plasticity observed in the external senses. Further, DMPFC modulation of IA networks may be an important mechanism by which MT alters information processing in the brain, increasing the contribution of interoception to perceptual experience. by gest on M ay 3, 2016 http://scaordjournals.org/ D ow nladed from Training interoceptive cortex Page 3 The psychologist Abraham Maslow famously argued that human behavior is motivated by a hierarchy of needs, ranking the body’s physiological requirements ahead of more abstract goals such as freedom, companionship, or social status (Maslow, 1943). Maslow’s hierarchy points to the complex interplay between two distinct representational systems for human attention: in cases of physiological imbalance, interoceptive attention (IA) is recruited to alert the individual to the body’s internal requirements (Liotti et al., 2001). When these needs are met, exteroceptive attention (EA) fosters exploratory behavior which aids in the pursuit of more conceptual goals (Gibson, 1988). In many cases, this transition from IA to EA is a natural and adaptive part of development, leading to a balanced sense of well-being as an individual becomes integrated with the social world (Ryan & Deci, 2000). However, placing a high importance on external goals can be problematic in the face of failure (Moberly & Watkins, 2010), as these events become diagnostic of an individual’s sense of self worth. In such situations, it may be difficult to disengage from patterns of negative cognitive elaboration that have become automatic and seemingly obligatory (Joorman & Siemer, 2011). Mindfulness training (MT) may be one means by which to alter the relationship between external events and self-attribution, limiting automatic self-evaluative processing (Frewen et al., 2008). MT often begins through courses such as Mindfulness-Based Stress Reduction (MBSR) (Kabat-Zinn, 1990), in which individuals learn meditation techniques in a weekly group setting, and then practice at home with guided meditation and yoga audio recordings. There have been several accounts of the cognitive mechanisms by which MT promotes salutary effects, such as decentering of experience (Fresco et al., 2007), a broadened context for appraisal (Garland et al., 2011), or otherwise “reperceiving” the world (Carmody et al., 2009). Despite progress in refining by gest on M ay 3, 2016 http://scaordjournals.org/ D ow nladed from Training interoceptive cortex Page 4 these cognitive models explaining higher-level effects of MT, we lack a translational account for how mindfulness practices directly modulate attention networks to promote cognitive change. We propose that the development of IA may be one foundation by which MT promotes cognitive change. Many mindfulness practices involve sustained attention to interoceptive sensations of respiration or bodily sensation, designed to improve the stability and frequency with which one perceives the transitory nature of human experience (Baer et al., 2006; Brown et al., 2007; Ivanovski & Malhi, 2007; Kabat-Zinn, 1982). While some exercises in MBSR investigate emotional reactivity to external events, relying on EA to recognize one’s behavioral patterns in the world, most MBSR practices employ IA, cultivating sustained attention towards bodily sensation in response to stress. It may be this ability to skillfully recruit IA that disrupts automatic conceptual elaboration and allows for more adaptive regulatory strategies to be invoked. Supporting this notion, in prior work we have demonstrated that MBSR reduces the involuntary recruitment of a cognitive elaboration network, instead promoting recruitment of viscerosomatic regions associated with momentary awareness of internal sensation (Farb et al., 2007), and that this improved access to body sensation during sadness is associated with lower levels of depression (Farb et al., 2010). In the present study, we investigated whether practicing sustained IA as through an MBSR course modulates neural representation networks for