1 s2.0 S0149763415302244 Main

Neuroscience and Biobehavioral Reviews 65 (2016) 208–228
Contents lists available at ScienceDirect
Neuroscience and Biobehavioral Reviews

journal homepage: www.elsevier.com/locate/neubiorev
Review article
Functional neuroanatomy of meditation: A review and meta-analysis

of 78 functional neuroimaging investigations
Kieran C.R. Fox a,∗ , Matthew L. Dixon a , Savannah Nijeboer a , Manesh Girn a ,
James L. Floman b , Michael Lifshitz c , Melissa Ellamil d , Peter Sedlmeier e ,
Kalina Christoff a,f
a
Department of Psychology, University of British Columbia, 2136 West Mall, Vancouver, B.C., V6T 1Z4, Canada
b
Department of Educational and Counselling Psychology, and Special Education, University of British Columbia, 2125 Main Mall, Vancouver, B.C., V6T 1Z4,
Canada
c
Integrated Program in Neuroscience, McGill University, 3775 University St., Montreal, QC, H3A 2B4, Canada
d
Neuroanatomy and Connectivity Research Group, Max Planck Institute for Human Cognitive and Brain Sciences, Stephanstrasse 1a, Leipzig, 04103,
Germany
e
Institut für Psychologie, Technische Universität Chemnitz, 43 Wilhelm-Raabe Street, Chemnitz, Germany
f
Brain Research Centre, University of British Columbia, 2211 Wesbrook Mall, Vancouver, B.C., V6T 2B5, Canada
a r t i c l e i n f o a b s t r a c t
Article history: Meditation is a family of mental practices that encompasses a wide array of techniques employing
Received 7 October 2015 distinctive mental strategies. We systematically reviewed 78 functional neuroimaging (fMRI and PET)
Received in revised form 25 February 2016 studies of meditation, and used activation likelihood estimation to meta-analyze 257 peak foci from 31
Accepted 1 March 2016
experiments involving 527 participants. We found reliably dissociable patterns of brain activation and
Available online 28 March 2016
deactivation for four common styles of meditation (focused attention, mantra recitation, open mon-
itoring, and compassion/loving-kindness), and suggestive differences for three others (visualization,
Keywords:
sense-withdrawal, and non-dual awareness practices). Overall, dissociable activation patterns are con-
Meditation
Open monitoring gruent with the psychological and behavioral aims of each practice. Some brain areas are recruited
Focused attention consistently across multiple techniques—including insula, pre/supplementary motor cortices, dorsal
Functional magnetic resonance imaging anterior cingulate cortex, and frontopolar cortex—but convergence is the exception rather than the
Meta-analysis rule. A preliminary effect-size meta-analysis found medium effects for both activations (d = 0.59) and
Activation likelihood estimation deactivations (d = −0.74), suggesting potential practical significance. Our meta-analysis supports the neu-
rophysiological dissociability of meditation practices, but also raises many methodological concerns and
suggests avenues for future research.
© 2016 Elsevier Ltd. All rights reserved.
Contents
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
1.1. Functional neuroimaging of meditation: the need for quantitative meta-analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
1.2. Four general categories of meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
1.2.1. Focused attention meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
1.2.2. Mantra recitation meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
1.2.3. Open monitoring meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
1.2.4. Loving-kindness and compassion meditations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
1.3. Other forms of meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
1.4. Delineating reliable neural correlates of different meditation practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
1.5. Are the effects of meditation practices on brain function of any practical significance? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
∗ Corresponding author.
E-mail address: [email protected] (K.C.R. Fox).
http://dx.doi.org/10.1016/j.neubiorev.2016.03.021
0149-7634/© 2016 Elsevier Ltd. All rights reserved.
K.C.R. Fox et al. / Neuroscience and Biobehavioral Reviews 65 (2016) 208–228 209
2. Meta-analytic methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213

2.1. Literature review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
2.1.1. Search strategy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
2.1.2. Study inclusion and exclusion criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.1.3. Study classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.2. The question of baselines, control conditions, and tasks engaged in during meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.3. Reporting and classification of results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.4. Activation likelihood estimation (ALE) meta-analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.4.1. General methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.4.2. Primary neuroimaging meta-analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
2.5. Effect size meta-analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
2.6. Estimating publication bias in meta-analytic results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
3. Results I: neuroimaging meta-analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
3.1. Focused attention meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
3.2. Mantra recitation meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
3.3. Open monitoring meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
3.4. Loving-kindness and compassion meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
4. Results II: effect size meta-analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
5. Discussion, future directions, and conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
5.1. Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
5.2. Functional neuroanatomy of focused attention meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
5.2.1. Activations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
5.2.2. Deactivations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.3. Functional neuroanatomy of mantra recitation meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.3.1. Activations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.3.2. Deactivations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.4. Functional neuroanatomy of open monitoring meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.4.1. Activations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.4.2. Deactivations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.5. Functional neuroanatomy of loving-kindness and compassion meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.5.1. Activations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
5.5.2. Deactivations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.6. Convergent findings across meditation categories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.6.1. Insular cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.6.2. Premotor cortex and supplementary motor area (lateral/medial BA 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.6.3. Dorsal anterior/mid cingulate cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.6.4. Frontopolar cortex/rostrolateral prefrontal cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
5.7. Other meditation practices: further evidence for dissociability at the neural level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
5.8. Are contemplative practices dissociable by electrophysiological and neurochemical measures? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
5.9. Do functional neural effects associated with meditation have practical significance? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
5.10. Meditation, the brain, and clinical disorders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
5.11. Meta-analytic limitations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
5.12. Other methodological challenges in the neuroimaging of meditation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
5.12.1. Lifestyle variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
5.12.2. Pre-existing differences in brain structure and function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
5.12.3. ‘Overshadowing’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
5.12.4. Enduring trait differences in brain and cognitive function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
5.12.5. Integration of functional neuroimaging methods with behavioral measures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
5.13. Summary and conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Appendix A. Supplementary data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
1. Introduction and mounting empirical evidence suggests that meditation indeed

leads to or is associated with significant changes in cognitive and
Meditation has been used as a tool to train the mind for thou- affective processing (Sedlmeier et al., 2012), as well as alterations
sands of years (Analayo, 2003; Iyengar, 2005). Broadly speaking, in brain structure (Fox et al., 2014) and function (Cahn and Polich,
meditation practices involve the monitoring and regulation of 2006).
attention and emotion (Lutz et al., 2008b; Tang et al., 2015). Med- Although meditative practices now garner serious interest
itation can be either directed outward to particular objects and from the cognitive neuroscience community (Fig. 1), meditation
sensory stimuli or turned inward to the workings of the mind and still tends to be viewed as a somewhat uniform practice (e.g.,
felt experiences of the body. A common principle across all forms of Sperduti et al., 2012). The umbrella-term ‘meditation’ encom-
meditation is that through specific and regular practice, the ability passes a wide variety of distinct practices with specific goals
to monitor and regulate mental-physical processes can be progres- and methods, however. The varying methods and scope of par-
sively developed—the way performance on any other cognitive or ticular forms of meditation include focalizing and sustaining
motor skill can be improved with practice (Ericsson et al., 1993; attention (Wallace, 2006), generating and maintaining complex
Slagter et al., 2011; Vago, 2014). Interest in the neural basis of med- visual imagery (Kozhevnikov et al., 2009), improving emotion regu-
itative practices has increased enormously over the past decade,
210 K.C.R. Fox et al. / Neuroscience and Biobehavioral Reviews 65 (2016) 208–228
synthesis, we sought to build on prior reviews and meta-analyses

by examining the possibility that comparable dissociations might
exist with modalities that measure cerebral blood flow (positron
emission tomography; PET) or blood-oxygenation level (functional
magnetic resonance imaging; fMRI). Other modalities, such as EEG,
have extremely high temporal resolution but very poor spatial res-
olution (or vice versa, in morphometric neuroimaging). Functional
neuroimaging methods, on the other hand, are unique in their
combination of good temporal and spatial resolution. This allows
for precise localization of dynamic changes in brain activity, pro-
viding unified spatiotemporal neurophysiological data that could
prove critical in differentiating one form of meditation practice
from another.
Taken together, our central aims were to: (i) comprehensively
review and meta-analyze the existing functional neuroimaging
studies of meditation (using the meta-analytic method known as
activation likelihood estimation, or ALE), and compare consisten-
cies in brain activation and deactivation both within and across
psychologically distinct meditation techniques; (ii) examine the
Fig. 1. The rapid increase in functional (fMRI and PET) and morphometric neu-
magnitude of the effects that characterize these activation pat-
roimaging of meditation and related secular contemplative practices. Functional terns, and address whether they suggest any practical significance;
neuroimaging (fMRI and PET) study counts based on the results of the present and (iii) articulate the various methodological challenges facing the
review; morphometric study counts based on our previous meta-analysis of this emerging field of contemplative neuroscience (Caspi and Burleson,
literature (Fox et al., 2014).
2005; Thompson, 2009; Davidson, 2010; Davidson and Kaszniak,
2015), particularly with respect to functional neuroimaging studies
lation and well-being (Chambers et al., 2009; Sedlmeier et al., 2012), of meditation.
and deepening compassion for others (Galante et al., 2014).
Despite the diversity of meditation practices, few behavioral,
clinical, or neuroimaging studies have directly compared medita- 1.1. Functional neuroimaging of meditation: the need for
tion styles within or across studies. As such, little is known about quantitative meta-analysis
whether functional neural differences underlie each type of prac-
tice (although see Lou et al., 1999; Manna et al., 2010; Brewer There are now close to a hundred functional neuroimaging stud-
et al., 2011; Lee et al., 2012; Tomasino et al., 2013). Although ies of meditation (Fig. 1)—but can any conclusions be drawn from
mounting empirical evidence suggests that these distinctive psy- this large and fast growing literature? Determining reliable neu-
chological practices may be dissociable at the neurophysiological ral correlates of any mental process is a difficult task under the
level (Lutz et al., 2008b; Travis and Shear, 2010a; Josipovic et al., best of conditions. This is all the more true when the mental pro-
2011; Tomasino et al., 2013), no comprehensive overview has ade- cesses in question are subtle and largely subjective, and when
quately addressed this complex issue. We therefore attempted to the participants employed are highly experienced practitioners
synthesize the large and rapidly expanding (Fig. 1) body of work on often operating within spiritual-religious frameworks. Despite the
the functional neuroanatomy of meditation practices in the present voluminous functional neuroimaging literature on meditation, we
review and meta-analysis. suspect that most researchers would be hard-pressed to provide a
Our central hypothesis was a simple one: meditation practices clear answer to the ostensibly simple question: What brain regions
distinct at the psychological level () may be accompanied by dis- are reliably recruited, for instance, by focused attention medita-
sociable activation patterns at the neurophysiological level (). tion? We expect that similar challenges would face researchers
Such a model describes a ‘one-to-many’ isomorphism between seeking to describe other common categories of meditation. Part of
mind and brain: a particular psychological state or process is the problem is that despite the rapid proliferation of neuroimaging
expected to have many neurophysiological correlates from which, studies of meditation in the past few years (Fig. 1), neither narrative
ideally, a consistent pattern can be discerned (Cacioppo and (Rubia, 2009; Tang et al., 2015), nor meta-analytic (Sedlmeier et al.,
Tassinary, 1990). 2012; Fox et al., 2014), nor theoretical (Holzel et al., 2011; Vago
Supporting or disconfirming this prediction through empirical and Silbersweig, 2012) synthesis of these results has been able to
synthesis is challenging, however, because at present there are few keep pace with the emergence of new and complex data. Given
(if any) ‘objective’ third-person measures that can confirm engage- the numerous and often contradictory results reported to date,
ment in a particular subjective state or mental practice (Lutz and researchers interested in the neural correlates of meditation may
Thompson, 2003; Fazelpour and Thompson, 2015). Current inves- find it difficult to decide among different findings. As experts on
tigations therefore necessarily rely on first-person reports that a meta-analytic methodology recently framed the general problem
specific mental practice is indeed being carried out as instructed or of integrating findings:
intended. Acknowledging this difficulty, we reasoned that if medi- “How should scientists proceed when results differ? First, it is
tation practitioners are really engaging in the practices they report clear how they should not proceed: they should not pretend that
to be—and if these practices engender their intended effects and there is no problem, or decide that just one study (perhaps the most
mental states—then distinctive patterns of brain activation and recent one, or the one they conducted, or a study chosen via some
deactivation should accompany each technique. other equally arbitrary criterion) produced the correct finding. If
In line with our hypothesis, it has been argued that various results that are expected to be very similar show variability, the
categories of meditation are characterized by differential elec- scientific instinct should be to account for the variability by further
troencephalography (EEG) signatures (Travis and Shear, 2010a), systematic work (Cooper and Hedges, 2009; p. 4)”.
cognitive-emotional effects (Sedlmeier et al., 2012), and poten- It is in this spirit of systematic, quantitative synthesis that the
tially neuroanatomical alterations (Fox et al., 2014). In the present present meta-analysis were conducted.
Although the coordinate-based meta-analysis we employed out being distracted appears to become progressively enhanced
here (see Section 2) is far from the perfect (or only) method of (e.g., MacLean et al., 2010) and less effortful (Wallace, 1999, 2006;
achieving this kind of synthesis (Salimi-Khorshidi et al., 2009), Brefczynski-Lewis et al., 2007). In particular, focused attention
it is a major improvement over narrative (and/or selective) sur- meditation may lead to three types of improvements: monitoring
veys of the literature, which often lead to erroneous conclusions the locus of attention, disengaging from distraction, and shifting
(Schmidt, 1992). We therefore echo other neuroscientists in argu- attention back to its intended target (Lutz et al., 2008b).
ing that quantitative meta-analysis of neuroimaging data is a potent
(indeed necessary) tool for better understanding the neural basis 1.2.2. Mantra recitation meditation
of mental processes (Wager et al., 2007, 2009; Salimi-Khorshidi Focused attention meditation itself may be subdivided into dis-
et al., 2009; Kober and Wager, 2010; Yarkoni et al., 2010; Hupé, tinct practices, or distinguished from practices that appear similar
2015). More specifically, meta-analysis can provide: (i) a less- at first glance: in particular, mantra recitation meditation involves
biased overview of an evidence base than a simple narrative review unique components (Travis, 2014). Mantra meditation—probably
or qualitative survey of prior work (Schmidt, 1992); (ii) specific best known in the form of the widespread ‘Transcendental
peaks of meta-analytic activation, rather than just broad regional Meditation’—involves the repetition of a sound, word, or sentence
results; (iii) statistically significant activation overlaps across stud- (spoken aloud or silently in one’s head) with the goals of calming
ies, instead of merely indicating broad ‘replications’; and (iv) a way the mind, maintaining focus, and avoiding mind-wandering. While
to mitigate the lack of statistical power caused by small samples mantra meditation therefore clearly overlaps with other forms of
found in many neuroimaging studies—a problem especially preva- focused attention in regard to its aims, it differs in that the object
lent in the study of meditation practitioners, where experienced of focus is a voluntary verbal-motor production, rather than natu-
individuals are difficult to recruit. For a deeper discussion of the rally arising body sensations (like the breath) or external physical
limitations of these meta-analytic methods, however, see Section objects (such as a point in space upon which the gaze is focused).
5.11. Further, mantra meditation appears to be associated with neural
Moreover, persuasive arguments have also been made for a correlates separate from other related forms of focused attention
move toward a more ‘cumulative’ science that focuses on broad practice (e.g., Lazar et al., 2000; Shimomura et al., 2008; Davanger
conclusions drawn from dozens or hundreds of independent inves- et al., 2010; Tomasino et al., 2013)—although, in the absence of a
tigations, as opposed to ‘critical experiments’ from isolated but meta-analysis, these differences remain only suggestive. Although
influential studies (Schmidt, 1992; Yarkoni et al., 2010). Some mantra meditation could reasonably be placed within the general
statisticians have gone as far as claiming that “any individual study ‘focused attention’ category of practices, because of its unique ele-
must be considered only a single data point to be contributed to a ments (most notably its linguistic, verbal-motor component) we
future meta-analysis” (Schmidt, 1992). Without endorsing such an examined it separately here.
extreme viewpoint, we agree that a careful sifting and quantitative
synthesis of the research conducted to date will help to advance 1.2.3. Open monitoring meditation
the state of the field. Open monitoring practice typically begins with bringing atten-
tion to the present moment and impartially observing all mental
1.2. Four general categories of meditation contents (thoughts, emotions, sensations, etc.) as they naturally
arise and subside. A key element of this practice is possessing
Before conducting a synthesis of the meditation neuroimaging an open, accepting attitude toward, and learning to ‘let go’ of,
literature, one must ask a central question: How are medita- mental content—neither resisting nor elaborating upon anything
tion practices to be categorized? There is growing consensus that that surfaces in awareness (Harvey, 1990; Walker, 1995; Suzuki,
at least three broad categories of meditation techniques can be 2003; Kabat-Zinn, 2005; Wangyal and Turner, 2011). In contrast to
discerned: focused attention, open monitoring, and compassion or focused attention, then, in open monitoring meditation mental con-
loving-kindness practices (Cahn and Polich, 2006; Lutz et al., 2008b; tent is neither evaluated for relevance to a particular goal, nor is any
Brewer et al., 2011; Vago and Silbersweig, 2012; Lippelt et al., 2014). content suppressed. This ‘non-elaborative’ mental stance cultivates
Similar classification schemes appear to reach back centuries to a more present-centered awareness in tune with the moment-to-
early Eastern treatises on meditation (Harvey, 1990; Lodro, 1998; moment experience of the mind and body (Analayo, 2003; Farb
Goenka, 2000; Wangyal and Turner, 2011). Although a few broad et al., 2007). Somatic and visceral body sensations are often a
categories do not begin to exhaust the subtleties of all contempla- prominent feature of this present moment experience, and hence
tive practices (Niranjananada, 1993; Singh, 2002), there seems to be interoceptive and exteroceptive inputs generally receive greater
fair justification for a tentative division along these lines (Lutz et al., processing in open monitoring than in focused attention practices
2008b; Vago and Silbersweig, 2012). The main aim of this review (e.g., Fox et al., 2012; Kerr et al., 2013). Open monitoring meditation
and meta-analysis was to investigate whether some isomorphism can also sometimes serve as a platform from which practitioners
(even if a complex one) can be discovered between psychological can go on to enter subtler states of non-dual awareness (see Section
practices and brain recruitment (Cacioppo and Tassinary, 1990): 1.3, below).
that is, are these various categories of meditation, which differ
markedly at the psychological level (), in fact characterized by 1.2.4. Loving-kindness and compassion meditations
distinctive patterns of brain activation and deactivation ()? Loving-kindness meditation (closely related to, but not iden-
tical with, compassion meditation) aims to deepen feelings of
1.2.1. Focused attention meditation sympathetic joy for all living beings, as well as promote altruis-
Focused attention meditation involves directing attention to one tic behaviors (Harvey, 1990; Gyatso and Jinpa, 1995; Kabat-Zinn,
specific object (e.g., the breath or a mantra) while monitoring and 2005; Lutz et al., 2008a. Typically, practitioners begin by generating
disengaging from extraneous thoughts or stimuli (Harvey, 1990; feelings of kindness, love, and joy toward themselves, then progres-
Hanh, 1991; Kabat-Zinn, 2005; Lutz et al., 2008b; Wangyal and sively extend these feelings to imagined loved ones, acquaintances,
Turner, 2011). Attention is directed to a particular object (most strangers, enemies, and eventually all living beings (Harvey, 1990;
commonly the sensations associated with respiration), and when- Kabat-Zinn, 2005; Lutz et al., 2008a). Compassion meditation gen-
ever the mind wanders, attention is redirected to this target. With erally takes this practice a step further: practitioners imagine the
regular training, the ability to voluntarily control attention with- physical and/or psychological suffering of others (ranging from
loved ones to all humanity) and cultivate compassionate attitudes detachment from or blocking of sensory inputs, including pain,
and responses to this suffering. Both practices share the long-term with the eventual goal of transcending any narrow sense of self
goals of enhancing sympathetic joy and empathy for pain, which are or personal identity. Although this type of practice shares many
viewed as trainable skills that increase altruistic behavior (Gyatso features of focused attention techniques (e.g., a highly selective
and Jinpa, 1995; Lutz et al., 2008a). Although there are potentially attentional focus), and would traditionally be practiced alongside
important psychological differences between these practices, of the them (Iyengar, 2005), it could also be considered a practice in its
neuroimaging studies to date, the instructions have been compa- own right. However, so far only one study has investigated neural
rable: participants have been asked to generate positive emotions correlates of this kind of technique (Kakigi et al., 2005).
(such as loving-kindness, well-wishing, or compassion) directed Non-dual awareness practices are another example. In general,
toward others and/or generated in a non-referential way (see the the aim of these practices is to dissolve or attenuate the bound-
Supplementary material for the exact instructions in each study). ary between subject and object (Josipovic, 2010; Travis and Shear,
We therefore grouped the available studies together, reasoning 2010b; Dunne, 2011). Although this kind of practice bears some
that they were more similar than different: despite some differ- resemblance to open monitoring meditation, it could be consid-
ence in instructions and implementation of the practices, all studies ered a distinct practice, and further investigation into its neural
involved a strong focus on cultivating positive affect directed out- correlates would be a welcome follow-up to a recent, seminal study
ward. We acknowledge, however, that differences in the various (Josipovic et al., 2011; Josipovic, 2014).
stages of these practices (e.g., compassion for self, vs. a loved one, Finally, another suite of practices aims at manipulating or
vs. a stranger, vs. non-referential compassion) might very well enhancing meta-awareness during various stages of sleep and
be resolved at the neural level by future research. Note, too, that dreaming. Yoga nidra (literally ‘sleep yoga’), for instance, involves
the instructions involved in the studies investigated here do not the deliberate dampening of sensory inputs to induce a ‘hypna-
exhaust the possibility of this type of practice, which can also gogic’ state (Mavromatis, 1987; Hori et al., 1994; Hayashi et al.,
involve complex visualizations and a focus on spiritual ‘benefac- 1999; Stenstrom et al., 2012), so as to enhance one’s capacity to
tors’ (Makransky et al., 2012). Further research is clearly warranted carry out visualization and relaxation practices (Saraswati, 1984).
to more thoroughly examine the many variants of loving-kindness Dream yoga (rmi-lam in Tibetan; svapnadarsana in Sanskrit) prac-
and compassion meditation. tices aim at enhancing meta-awareness during dreaming (i.e.,
In one sense, loving-kindness and compassion meditations enhancing so-called ‘lucid’ dreaming; Gackenbach and LaBerge,
could also be considered a form of focused attention in that they 1988), and are particularly prevalent in the Tibetan Buddhist tradi-
often focus intensively on a single object (the person who is the tion (Norbu and Katz, 1992; Mullin, 1996; Rinpoche, 2004). Other
target of the loving-kindness) and cultivate a consistent emotional techniques even aim at maintaining awareness during states of
tone (to the exclusion of other kinds of affect). Alternatively, when deep, dreamless sleep (Aurobindo, 2004; Sharma, 2012). We are
conducted in a non-referential, all-embracing way (extending one’s aware of only a single study of these practices, however (Kjaer et al.,
compassion and joyful feelings to any being or object that arises in 2002).
consciousness indiscriminately, or even cultivating such feelings
without any object whatsoever), these practices could conceivably
be considered a form of open monitoring. Nonetheless, given the 1.4. Delineating reliable neural correlates of different meditation
strong emphasis on deliberate cultivation of joyful, altruistic, and practices
empathetic emotions that is fairly unique to these practices, there
seems sufficient justification for considering them as a separate Only in about the past year have a sufficient number of studies
class, worthy of investigation in its own right. been cumulatively reported to allow for a reliable meta-analysis
of each major meditation type (including open monitoring and
1.3. Other forms of meditation loving-kindness/compassion meditations). Even very recent meta-
analyses have been forced by a paucity of empirical data to examine
It is important to note that these four putative categories of med- only focused attention and mantra recitation meditation (Tomasino
itation are intended not as a final classification scheme, but merely et al., 2013), or to group all practice types together (Sperduti et al.,
as a first step toward delineating quantitative neurophysiological 2012). Other meta-analyses have employed less theoretically or
() correlates associated with relatively distinctive psychological empirically grounded categorizations, such as basing meta-analytic
() practices (Cacioppo and Tassinary, 1990). There are several contrasts on the religion of origin for a practice (Buddhism vs.
other major categories of meditation that have been investigated Hinduism) as opposed to the phenomenological content, intended
in some preliminary work, but that could not be examined meta- goals, and behavioral characteristics of the practice itself (Tomasino
analytically due to a paucity of data. We nonetheless sought to et al., 2014).
examine and compare our meta-analytic results with these data Here we present the first meta-analysis inclusive of four major
by conducting a qualitative review of the results reported to date. categories of meditation, with a focus on elucidating whether
Visualization meditation, for instance, involves creating and sus- ostensibly distinct forms of mental practice activate a common
taining complex mental imagery for extended periods of time. neural substrate (Sperduti et al., 2012) or instead show disso-
These visualizations typically involve mandalas or yantras (geo- ciable patterns of underlying neural activation and deactivation.
metric designs) or deities or patron-teachers (Gyatso, 1981), and Accordingly, we conducted four separate activation likelihood esti-
experts in these techniques have been shown to have enhanced mation (ALE) meta-analyses using 257 foci of peak activation from
visuospatial processing (Kozhevnikov et al., 2009). Nonetheless, 25 independent PET and fMRI studies involving 31 unique experi-
only a single study of which we are aware has investigated the ments (Tables 1 and 2) to examine the patterns of brain activation
neural correlates of any form of visualization meditation (Lou associated with focused attention, mantra recitation, open moni-
et al., 1999), although loving-kindness and compassion practices toring, and loving-kindness/compassion meditation. Additionally,
can also sometimes involve important visualization components we examined the results for three categories of meditation that
(Weng et al., 2013). have only been investigated thus far in a single study each (Table 1).
Another almost uninvestigated set of practices based in yogic Because this paucity of data precluded a formal, quantitative meta-
traditions involve the withdrawal of the senses (pratyahara in analysis, we qualitatively reviewed the results for each practice
Sanskrit; (Iyengar, 2005). These practices involve the deliberate type and compared it to our other meta-analytic results (Table 1).
Table 1 late to clinical significance or relevance (Rosnow and Rosenthal,

Seven meditation categories investigated via quantitative meta-analysis and quali-
1989).
tative review.
An additional benefit of calculating effect sizes is that they can be
Meditation Type Contributing Contributing Foci used to estimate the extent of publication bias in a field (Egger et al.,
Experiments (N) A/D (Total)
1997), a major and outstanding issue across the social and neuro-
Quantitative meta-analysis biological sciences, including contemplative neuroscience. Because
Focused attention 7 48/13 (61) we found strong evidence for publication bias in morphometric
Mantra recitation 8 63/19 (82)
(anatomical) neuroimaging studies of meditation in a recent review
Open monitoring 10 45/29 (74)
Compassion/Loving-kindness 6 36/4 (40) (Fox et al., 2014), we sought to address whether or not such a bias
Totals 31 192/65 (257) might be affecting functional neuroimaging studies of meditation
Qualitative review
as well. The second central aim of this meta-analytic synthesis was
Visualization 1 – therefore to provide preliminary estimates of effect sizes in the
Sense-withdrawal 1 – functional neuroimaging of meditation in order to assess practical
Non-dual awareness 1 – significance and the possibility of publication bias.
Note: Some studies examined multiple meditation types and therefore contributed
to multiple analyses. This resulted in a total of 31 experiments (contrasts) from the
25 independent studies included in the quantitative meta-analyses for the first four
categories (see Table 2 for details of included studies). Further, three other categories 2. Meta-analytic methods
of meditation were also examined where results have been reported from only a sin-
gle study to date. These investigations were not included in any formal, quantitative 2.1. Literature review
meta-analyses, but the results were nonetheless examined and compared with our
meta-analytic findings in a qualitative fashion. A = activations; D = deactivations.
2.1.1. Search strategy
Three authors (KCRF, SN, and MG) searched MEDLINE (http://
www.ncbi.nlm.nih.gov/pubmed/), Google Scholar (http://scholar.
1.5. Are the effects of meditation practices on brain function of google.com), and PsycINFO (http://www.apa.org/pub/databases/
any practical significance? psycinfo/index.aspx) for all papers containing the word ‘medita-
tion’ since the first functional neuroimaging study of contemplative
Determining whether there are any consistencies in brain acti- practices was published (Lou et al., 1999). These extensive lists
vation for various styles of meditation is only the first step. A of articles were then refined by searching within results for
logical next question is whether activations and deactivations studies that contained any of the words or phrases ‘magnetic res-
associated with a given meditation technique have any prac- onance imaging’, ‘MRI,’ ‘neuroimaging,’ or ‘brain’ within the title
tical significance. That is, what effect sizes characterize these or abstract. Of the remaining results, every abstract was consulted
differences, and do effect sizes vary across particular forms of med- to see whether the study actually employed functional neuroimag-
itation? Note, however, that practical significance is not a fully ing methods to study meditation. The reference lists of each study
objective measure-typically a half-standard deviation difference found, as well as those of several major reviews (e.g., Holzel et al.,
is considered a notable effect, but this boundary is essentially 2011; Vago and Silbersweig, 2012), were also consulted to ensure
arbitrary, and ‘practical’ significance does not necessarily trans- that no studies were missed.
Table 2
Studies included in the meta-analyses (n = 25).
Study Practice N (M/C) Tradition Expertise Average Experience
Lou et al. (1999) OM + LK 9/– Yoga Nidra LTP >5 years

Lazar et al. (2000) MR 5/− Kundalini LTP 4 years
Brefczynski-Lewis et al. (2007) FA 14/16 Tibetan Buddhist LTP 19000 h & 44000 h
Farb et al. (2007) OM 20/− MBSR STT ∼42 h
Lutz et al. (2008a) LK 16/16 Tibetan Buddhist LTP 45 ± 12.7 years
Shimomura et al. (2008) MR 8/− Pure Land Buddhism LTP >10 years
Lutz et al. (2009) LK 10/12 Tibetan Buddhist LTP 40 ± 9.6 years
Davanger et al. (2010) MR 4/− Acem Meditation LTP >23 years
Engström et al. (2010) MR 8/− Acem Meditation and Kundalini Yoga LTP 1.17 years
Manna et al. (2010) FA + OM 8/− Vipassana (Theravada—Thai Forest Tradition) LTP 15750 h
Brewer et al. (2011) FA + OM + LK 12/13 Vipassana (Mindfulness) LTP 10565 ± 5148 h
Ives-Deliperi et al. (2011) OM 10/− MBSR LTP >4 years
Kalyani et al. (2011) MR 12/− Om mantra recitation LTP + STT Not reported
Taylor et al. (2011) OM 12/10 Zen LTP + STT LTP: >1000 h STT: 1 wk
Wang et al. (2011) MR 10/− Kundalini yoga LTP ∼20,000 h
Dickenson et al. (2012) FA 31/− Mindfulness/MBSR STT No previous experience;
brief meditation
induction only
Gard et al. (2012) OM 17/17 Vipassana LTP 5979 ± 5114
Hasenkamp et al. (2012) FA 14/− Various LTP 1386 ± 1368 h
Lee et al. (2012) FA + LK 22/22 Theravada LTP FA: 5249 ± 6192 h
LK: 7492 ± 6681 h
Farb et al. (2013) OM 20/16 MBSR STT 49.2 ± 4.3 h
Guleria et al. (2013) MR 14/− Soham Meditation LTP 2088 ± 320 h
Lutz et al. (2013) OM 14/14 Tibetan Buddhist LTP 27000 ± 12500 h
Weng et al. (2013) LK 20/21 Tibetan Buddhist STT 5.9 ± 0.5 h
Lutz et al. (2014) OM 24/22 Mindfulness/MBSR STT Variable
Xu et al. (2014) FA + MR 14/− Acem Meditation LTP 27 ± 9 years
All studies employed fMRI except for Lou et al. (1999), which employed PET. C: controls; FA: focused attention; LK: loving-kindness or compassion meditation; LTP: long-term
practitioners; M: meditators; MBSR: mindfulness-based stress reduction; MR: mantra recitation; N: sample size; OM: open monitoring; STT: short-term training.
2.1.2. Study inclusion and exclusion criteria 2.2. The question of baselines, control conditions, and tasks
All studies using functional neuroimaging to investigate some engaged in during meditation
form of meditation were considered. By ‘functional neuroimaging’
we mean functional magnetic resonance imaging (fMRI) or positron One potential problem is that even studies examining a sim-
emission tomography (PET) studies that can provide details about ilar form of meditation might compare a meditation practice
the specific location of brain activations and deactivations. Using to very different baseline or comparison tasks and conditions.
the search strategy detailed above, a total of 78 studies served as However, collapsing across numerous different baselines or con-
the initial pool of data to be reviewed (see Table 2 for included trol conditions is a common (in fact, usually inevitable) practice
studies, and Table S1 for excluded studies). in meta-analyses of functional neuroimaging studies (Turkeltaub
In order to ensure the most rigorous results possible, strin- et al., 2002; Wager et al., 2003, 2007, 2009; Caspers et al., 2010;
gent inclusion criteria were applied. Studies were required to: (i) Yarkoni et al., 2010; Tomasino et al., 2013). It is by no means a
report specific peak foci of activation in either Talairach or Mon- problem limited to the meditation literature. Aside from the het-
treal Neurological Institute (MNI) space; (ii) include a reasonable erogeneity of conditions across studies, meditation is also often
sample size (i.e., case studies of single subjects were excluded); investigated during a given task or form of stimulus presentation.
(iii) involve participants who actually practiced meditation, be they Although typically these tasks or stimuli are equally present in the
novices or long-term practitioners (as opposed to, for example, baseline or control condition (and therefore, in principle, should
studies involving only self-report scales of ‘dispositional’ mindful- not unduly influence the meditation state results), the potential
ness); (iv) involve actual meditation during the scanning session (as for our results representing interactions between meditation and
opposed to having meditation practitioners engage in other tasks the given task employed should be kept in mind. These issues are
without some explicit contemplative element); and (v) involve discussed further in Section 5.
healthy, non-clinical populations. Finally, (vi) only reports pub- The goal of the present synthesis was to determine, irrespective
lished in peer-reviewed scientific journals were included (results of any incidental differences in comparison or baseline conditions
from conference abstracts, presented talks, dissertations, etc., were and tasks: (i) whether differing meditation practices tend to reliably
excluded). For further details on excluded studies, see the Supple- recruit differentiable neural networks; (ii) if the differences in brain
mentary material methods. activation show practically significant effect sizes; and lastly, (iii)
Ultimately, about one third (25 of 78) of the examined stud- whether distinct practices exhibit differing mean effect sizes (i.e.,
ies were included in our meta-analyses (Table 2). Given that some is a given style of meditation associated with greater differences in
of these studies examined multiple categories of meditation, we brain activity than others?).
investigated a total of 31 separate ‘experiments’ or contrasts of
interest spread across the four major meditation categories.
2.3. Reporting and classification of results
All peak voxel coordinates are reported in Montreal Neurolog-

2.1.3. Study classification ical Institute (MNI) space. For consistency, for all studies where
Two coders with longtime personal meditation experience peak voxels were originally reported in Talairach coordinates, we
(authors KCRF and MLD) independently classified all included used the WFU Pickatlas software package (Maldjian et al., 2003)
results as involving focused attention, open monitoring, compassion to perform a nonlinear transformation of Talairach coordinates
or loving-kindness, or mantra recitation meditation. Concordance to MNI space. For meta-analytic ALE results, region classifications
was nearly perfect: a single disparity was resolved after fur- follow those indicated in the Multi-Image Analysis GUI (‘Mango’)
ther discussion. Several studies employed multiple practice types image-viewing software (UT Health Science Center Research Imag-
across different sessions; in these cases, results from each session ing Institute) used to display our findings (see Results below). For
were coded separately and considered individual ‘experiments.’ additional precision, the Duvernoy neuroanatomical atlas was also
Therefore, the number of ‘experiments’ exceeded the number of consulted (Duvernoy et al., 1991).
independent studies actually included in the meta-analysis (see
Tables 1 and 2). All classifications for included studies are sum-
marized in Table 2. 2.4. Activation likelihood estimation (ALE) meta-analysis
Further, one study each was found that examined visualization
(Lou et al., 1999), sense-withdrawal (Kakigi et al., 2005), and non- 2.4.1. General methods
dual awareness (Josipovic et al., 2011) forms of meditation. Again, We used a quantitative, random-effects meta-analytic method
both coders (KCRF and MLD) independently classified and agreed known as activation likelihood estimation (ALE; Turkeltaub et al.,
on these categorizations. Due to there being only a single study for 2002, 2012; Laird et al., 2005; Eickhoff et al., 2009, 2012) imple-
each category, these reports were not included in any formal meta- mented in the software program GingerALE 2.1.1 (San Antonio,
analyses. Nevertheless, their results were qualitatively reviewed TX: UT Health Science Center Research Imaging Institute). The
and considered in comparison to our meta-analytic results for the most recent ALE algorithm tests for above-chance clustering of
other four general meditation styles (see Table 1). peak foci from different experiments included in the meta-analysis
In virtually all cases, classification was straightforward and mir- (Eickhoff et al., 2009, 2012) by comparing actual activation foci
rored the classifications described by the authors of the studies locations/clustering with a null distribution that includes the same
in their methods. For instance, phrases such as ‘concentration number of peak foci distributed randomly throughout the brain.
meditation’ were categorized under the focused attention cate- Included activation foci were smoothed using a full-width half
gory; mantra meditation studies were easily categorized based maximum (FWHM) Gaussian kernel dependent on the total sam-
on descriptions of mantra recitation in the methods; and so on. ple size of the experiment from which foci were drawn (larger
Because of the prevalence of classification schemes involving the sample → smaller smoothing kernel—empirically determined by
four categories described above (e.g., Cahn and Polich, 2006; Lutz Eickhoff et al., 2009, 2012). Resulting statistical maps show clus-
et al., 2008b; Travis and Shear, 2010a; Vago and Silbersweig, 2012), ters where convergence between foci of activation or deactivation
many studies in fact explicitly described their methods using these is greater than would be expected by chance (i.e., if foci from each
same (or highly similar) terms. experiment were distributed independently).
2.4.2. Primary neuroimaging meta-analyses roimaging data, so long as these serious limitations are noted and
We meta-analyzed a total of 257 peak foci of activation or attempts are made to correct for them. The many relevant issues
deactivation drawn from 25 studies (Table 2), involving 31 sep- are discussed at length in the Supplementary methods section.
arate ‘experiments’ or contrasts (summarized in Table 1). In Briefly, our procedure for applying these corrections was as
order to retain a maximal amount of information, this primary follows: After calculating effect sizes directly from the peak or
meta-analysis collapsed data from between-group (long-term maximum t or F statistics reported in the original studies, we then
practitioners vs. novices or controls) and repeated-measures (the adjusted these values downward to approximate more conserva-
same practitioners pre- and post-meditation training) designs. tive estimates of t or F statistics for the entire cluster of significant
Although only a small amount of data came from short-term difference. Next, we further deflated these values to account for
training (STT) investigations (6 of 25 studies included in the the inflationary bias of effect sizes derived only from results that
meta-analysis), nonetheless this difference presents a potentially have exceeded stringent statistical thresholds (Schmidt, 1992). In
confounding factor. We therefore addressed this issue by perform- these ways, we aimed to provide effect size estimates that more
ing supplemental meta-analyses that included only investigations closely approximate true effect sizes. It should be noted, however,
of long-term practitioners (see Supplementary materials). that we neither claim nor expect that this procedure is ideal. It is
Several coordinate foci (n = 7) from the included studies fell clear that some deflation of reported values is necessary (Schmidt,
outside of the brain mask templates used with the GingerALE meta- 1992; Yarkoni, 2009; Yarkoni et al., 2010; Hupé, 2015), but the
analysis software (1 focus for focused attention activations; 4 foci best method and optimal degree of deflation remains unclear. As
for open monitoring activations; and 2 foci for loving-kindness such, our methods are a preliminary effort to produce effect sizes
activations). This is a normal occurrence when using exclusive that are more accurate than those produced by calculations solely
meta-analytic template masks. The total number of foci included based on t-statistics reported in the literature, but should not by
in the final meta-analyses was therefore 250. any means be considered a definitive approach. For full details, as
Statistical maps were thresholded using a false discovery rate well as specific formulas used, see the Supplementary methods.
(FDR; Genovese et al., 2002) of q = 0.05 and a cluster threshold
of k = 100 mm3 . The meta-analytic software offers suggested min- 2.6. Estimating publication bias in meta-analytic results
imum cluster thresholds depending on the number of studies and
foci entered. These recommendations centered around 100 (gen- The bias toward publication of only positive (i.e., non-null)
erally, between 88 and 132 mm3 ), but differed slightly in each of results is a serious concern (the ‘file drawer’ problem; c.f. Rosenthal,
our eight meta-analyses (activations and deactivations for each of 1979). We constructed a funnel plot (scatterplot of effect size
four meditation categories). For consistency, we therefore set a against sample size) to test for potential publication bias in our
standard value of 100 mm3 for all analyses. Note that the cluster sample of studies (Egger et al., 1997). Effect sizes were calculated
threshold is essentially arbitrary and does not affect the underly- as described above and in the Supplementary methods, and plotted
ing meta-analytic analyses or results; it merely determines which against total sample size (meditators + controls). For detailed dis-
meta-analytic clusters are deemed ‘significant’ and therefore to be cussion of funnel plots see Egger and colleagues (1997). For other
reported in tables of results. It does not affect the underlying meta- examples of their use in meta-analyses of meditation and further
analytic results, which display all meta-analytic activations and discussion see Fox et al. (2014) and Sedlmeier et al. (2012).
deactivations regardless of cluster threshold. The full meta-analytic
data are visible in our supplementary results figures. 3. Results I: neuroimaging meta-analysis
To display our results, maps were first computed separately
for activations and deactivations for each meditation category. 3.1. Focused attention meditation
Activation and deactivation maps were then concatenated onto a
single template brain image for visualization purposes. The cre- Meta-analysis of focused attention studies resulted in 2 sig-
ation of the final figures for presentation here was performed nificant clusters of activation, both in prefrontal cortex (Table 3;
in the MRIcron software package using standard MNI brain Fig. 2). Activations were observed in regions associated with the
templates (http://www.mccauslandcenter.sc.edu/mricro/mricron/ voluntary regulation of thought and action, including the premo-
index.html) (Rorden et al., 2007). tor cortex (BA 6; Fig. 2b) and dorsal anterior cingulate cortex (BA
24; Fig. 2a). Slightly sub-threshold clusters were also observed in
2.5. Effect size meta-analysis the dorsolateral prefrontal cortex (BA 8/9; Fig. 2c) and left mid
insula (BA 13; Fig. 2e); we display these somewhat sub-threshold
In addition to determining which brain regions were consis- results here because of the obvious interest of these findings in
tently activated by various meditation practices, we sought to practices that involve top-down focusing of attention, typically
evaluate the magnitude of these differences (i.e., their effect sizes; focused on respiration. We also observed clusters of deactivation in
Cohen, 1992; Lipsey and Wilson, 2001; Sedlmeier et al., 2012). In regions associated with episodic memory and conceptual process-
all, 17 of the 25 studies that met our inclusion criteria also provided ing, including the ventral posterior cingulate cortex (BA 31; Fig. 2d)
necessary and sufficient data to allow reliable calculation of effect and left inferior parietal lobule (BA 39; Fig. 2f). A detailed series of
sizes. slices covering the entire brain is presented in Fig. S1.
Although the calculation and comparison of effect sizes in A supplementary meta-analysis, excluding studies that inves-
neuroimaging studies faces a number of challenges, simple null- tigated only practitioners with short-term training, yielded nearly
hypothesis significance tests and p-values have come under harsh identical results (see Table S2).
criticism as well (Schmidt, 1992; Schmidt and Hunter, 1997;
Cumming, 2012, 2013; Hupé, 2015). The meaning and relevance of 3.2. Mantra recitation meditation
null-hypothesis significance testing and associated statistics (such
as t and F statistics) has become so hotly contested that some jour- A meta-analysis of mantra recitation studies revealed seven sig-
nals have outright banned reporting of such values (Savalei and nificant activation clusters (Table 4). Activations were observed in
Dunn, 2015; Trafimow and Marks, 2015). regions associated with planning and executing voluntary motor
Bearing in mind these considerations, we concluded that effect output, including the posterior dorsolateral prefrontal cortex/left
sizes are worth calculating and reporting in meta-analyses of neu- premotor cortex (BA 6/8; Fig. 3d), pre-supplementary motor cortex,
Table 3
Activations and deactivations associated with focused attention meditation.
Region Cluster Size Side Peak Coordinates Peak ALE value

(mm3 ) (x, y, z)
Activations
Premotor cortex 712 L −36, 6, 56 (BA 6) 0.0144
Dorsal anterior/mid cingulate cortex 280 M 2, 12, 32 (BA 24) 0.0112
Deactivations
Posterior cingulate cortex 152 M −6, −60, 18 (BA 30) 0.0071
Inferior parietal lobule 144 L −48, −72, 30 (BA 39) 0.0071
Fig. 2. Peak activations and deactivations associated with focused attention meditation. (a) Dorsal anterior/mid cingulate cortex. (b) Premotor cortex/posterior dorsolateral
prefrontal cortex. (c) Sub-threshold cluster in dorsolateral prefrontal cortex. (d) Posterior cingulate cortex. (e) Sub-threshold cluster in mid-insula. (f) Inferior parietal lobule.
Warm colors: activations; cool colors: deactivations. See the Supplementary materials for a detailed series of slices covering the entire brain (Fig. S1).
Table 4
Activations and deactivations associated with mantra recitation meditation.
Region Cluster Size (mm3 ) Side Peak Coordinates (x, y, z) Peak ALE value
Activations
Premotor cortex 896 L −30,4, 60 (BA 6) 0.0137
Supplementary motor area 584 M 0, 12, 54 (BA 6) 0.0111
112 M −4, 2, 68 (BA 6) 0.0080
Putamen/Lateral globus pallidus 368 R 28, −16, −6 0.0111
Fusiform gyrus 248 R 40, −26, −30 (BA 20/36) 0.0093
Cuneus 160 R 24, −86, 26 (BA 18) 0.0082
Precuneus 152 L −14, −56, 54 (BA 7) 0.0082
Deactivations
Anterior insula 192 L −28, 25, −7 (BA 13) 0.0075
supplementary motor cortex (BA 6; Fig. 3a), and putamen/lateral identical results. The only notable difference was the disappearance
globus pallidus (Fig. 3b). Other activations were observed in regions of the single deactivation cluster in the anterior insula (see Table
associated with visual processing and mental imagery including S3). Although ALE implements controls for sample size, another
the fusiform gyrus (Fig. 3c), cuneus (BA 18), and precuneus (BA concern was the inclusion of two studies with very small sam-
7) (Fig. S2). There was also a small (but non-significant) cluster ple sizes—Lazar et al. (2000) with n = 5, and Davanger et al. (2010)
located within Broca’s area (BA 44/6), consistent with the verbal with n = 4–which might still have unduly influenced the results. We
component of mantra recitation. A single cluster of deactivation therefore also executed our analysis with data from these two stud-
was observed (Table 4) in the left anterior insula (BA 13)/claus- ies excluded. This had no effect on the pattern of deactivations. All
trum (Fig. 3b), a region associated with processing viscero-somatic remaining clusters of activation were the same, but two clusters
body signals. A detailed series of slices covering the entire brain is disappeared from the results: the putamen/lateral globus pal-
presented in Fig. S2. lidus, and the small (112 mm3 ) cluster in the supplementary motor
A supplementary meta-analysis, excluding studies that inves- area.
tigated only practitioners with short-term training, yielded nearly
Fig. 3. Peak activations and deactivations associated with mantra recitation meditation (a) Supplementary motor area. (b) Bilateral deactivations in the anterior insula (blue)
and activations in the right basal ganglia (putamen and globus pallidus). (c) Fusiform gyrus. (d) Premotor cortex. Warm colors: activations; cool colors: deactivations. See
the Supplementary materials for a detailed series of slices covering the entire brain (Fig. S2). (For interpretation of the references to colour in this figure legend, the reader is
referred to the web version of this article.)
3.3. Open monitoring meditation tivations were observed. A detailed series of slices covering the
entire brain is presented in Fig. S4.
Meta-analysis of open monitoring studies showed five signif- A supplementary meta-analysis, excluding studies that inves-
icant clusters of activation (Table 5). One cluster was located in tigated only practitioners with short-term training, yielded nearly
the insula (BA 13; Fig. 4d), consistent with awareness of ongo- identical results. The only notable difference was the appearance of
ing viscero-somatic body signals. Other significant clusters were a small cluster of activation in the left somatosensory cortices (see
observed in regions associated with the voluntary control of action, Table S5).
including the left inferior frontal gyrus (BA 44/45; Fig. 4b), pre-
supplementary motor area (BA 32/6; Fig. 4a), supplementary motor
4. Results II: effect size meta-analysis
area (BA 6; Fig. 4a and c), and premotor cortex (BA 6; Fig. 4c).
Smaller (non-significant) clusters were observed in the rostro-
As detailed in the Supplementary methods, we derived effect
lateral prefrontal cortex (BA 10) and mid-dorsolateral prefrontal
sizes (Cohen’s d) from studies that reported t or F statistics, and
cortex (BA 9/46)—regions associated with cognitive control and
then adjusted these raw effect sizes for the severe inflationary bias
metacognitive awareness. We observed a single significant cluster
that influences effect sizes in all neuroimaging research. From our
of deactivation (Table 5) in the right thalamus (Fig. 4d). A detailed
adjusted effect sizes for each individual reported result, we then
series of slices covering the entire brain is presented in Fig. S3.
calculated mean positive (+) and negative (−) effect sizes for each
A supplementary meta-analysis, excluding studies that investi-
study (and for each different practice within a study, if more than
gated only practitioners with short-term training, yielded similar
one was examined). These results are reported in Table 7. We then
results (Table S4). Some of the more salient differences are outlined
pooled these data to calculate mean (+) and (−) effect sizes for
in the Section 5, below (also, compare Table 5 with Table S4).
each type of practice (Table 8). Recall that in total, two deflation
coefficients were employed to adjust for suspected inflation bias.
Coefficients of 0.697 and 0.60, for a net coefficient of 0.4182, were
3.4. Loving-kindness and compassion meditation
applied to the effects calculated from the primary reports. There-
fore, in order to obtain the unadjusted values, simply multiply our
Meta-analysis of loving-kindness and compassion meditation
effect size results by 2.391. For details see Section 2.5.
revealed significant activation clusters in 3 regions (Table 6).
Activations were observed in regions associated with awareness
of bodily sensations and feelings including the right ante- 5. Discussion, future directions, and conclusions
rior insula/frontal operculum (BA 13; Fig. 5a) and secondary
somatosensory areas extending into the anterior inferior parietal 5.1. Overview
lobule (BA 2/40; Fig. 5b). Additionally, activation was observed near
the parieto-occipital sulcus (BA 23/31; Fig. 5c). No significant deac- The ensuing discussion is divided into three broad sections:
Table 5
Activations and deactivations associated with open monitoring meditation.
Activations
Supplementary motor area 824 M −6, 4, 60 (BA 6) 0.0165
Dorsal anterior cingulate cortex/pre-supplementary motor area 368 M −6, 18, 44 (BA 32/6) 0.0110
Insular cortex (mid/anterior) 360 L −44, 10, 4 (BA 13) 0.0111
Inferior frontal gyrus 200 L −50, 16, 14 (BA 44/45) 0.0092
Premotor cortex 104 L −44, 10, 46 (BA 6) 0.0089
Deactivations
Thalamus (pulvinar) 112 R 17, −24, 11 0.0084
Fig. 4. Peak activations and deactivations associated with open monitoring meditation. (a) Supplementary motor area (larger, more superior cluster) and dorsal anterior
cingulate cortex/pre-supplementary motor area. (b) Inferior frontal gyrus. (c) Premotor cortex (smaller cluster to left), and supplementary motor area also visible again
(center). (d) Activation in mid/anterior insula and bilateral deactivations in the thalamus. Warm colors: activations; cool colors: deactivations. L: left; R: right. See the
Supplementary materials for a detailed series of slices covering the entire brain (Fig. S3).
Table 6
Activations and deactivations associated with loving-kindness and compassion meditation.
Activations
Anterior insula 512 R 38, 22, 14 (BA 13) 0.0135
Parieto-occipital sulcus 344 R 24, −60, 18 (BA 23/31) 0.0115
Somatosensory cortices/Inferior parietal lobule 320 R 54, −26, 30 (BA 2/40) 0.0101
(i) First, we outline and interpret the various results from that may not have exceeded our cluster thresholds. We also inte-
our meta-analysis and qualitative review of neuroimaging find- grate the limited findings from the three meditation categories that
ings (Sections 5.2–5.8). We synthesize and simplify all of these could not be included in the quantitative meta-analysis, for the
results in a single summary figure (Fig. 7). We focus on the clus- same reasons: preliminary data, even if tentative, is still informa-
ters that attained statistical significance in our meta-analysis, but tive and relevant to our discussion. In a similar vein, the figures in
it should be recalled that significance and cluster thresholds are our Results section, for the sake of clarity, focus on the most reli-
ultimately arbitrary and should not restrict discussion. Some ‘sig- able, consistent clusters for each category of meditation. However,
nificant’ clusters are difficult to understand or explain, whereas we have presented detailed series of slices in our Supplementary
some non-significant clusters dovetail extremely well with the materials in order to clearly visualize all activations associated with
stated techniques and goals of a given form of meditation. We a given meditation type (not just those that exceeded our cluster
therefore endeavor to offer as integrated and impartial a discussion and significance thresholds; see Figs. S1–S4).
as possible, including consideration of meta-analytic activations
Fig. 5. Peak activations associated with loving-kindness and compassion meditation. (a) Right anterior insula/frontal operculum. (b) Right somatosensory cortices. (c)
Parieto-occipital sulcus. L: left; R: right. See the Supplementary materials for a detailed series of slices covering the entire brain (Fig. S4).
Table 7
Summary of adjusted mean effect sizes for each study and practice type.
Study Sample (N) Practice Mean (+) effect size (Cohen’s d) Mean (−) effect size (Cohen’s d)
Brefczynski-Lewis et al. (2007) 30 FA 0.51 –

Lutz et al. (2008a) 32 LK 0.33 –
Lutz et al. (2009) 22 LK 0.56 –
Davanger et al. (2010) 4 MR 1.48 –
Manna et al. (2010) 8 FA 1.56 −1.29
OM 1.25 –
Brewer et al. (2011) 25 FA – −0.71
OM – −0.58
LK – −0.68
Gard et al. (2011) 34 OM 0.46 –
Ives-Deliperi et al. (2011) 10 OM 2.03 −2.01
Kalyani et al. (2011) 12 MR – −1.11
Taylor et al. (2011) 22 OM – −0.79
Dickenson et al. (2012) 31 FA 0.49 –
Hasenkamp et al. (2012) 14 FA 0.67 –
Lee et al. (2012) 44 FA 0.56 –
LK 0.54 –
Farb et al. (2013) 36 OM 0.71 –
Lutz et al. (2013) 28 OM 0.58 −0.62
Weng et al. (2013) 41 LK 0.36 –
Lutz et al. (2014) 46 OM 0.51 −0.45
Table 8
Summary of adjusted mean effect sizes by meditation practice type.
Meditation Type Contributing Experiments (N) Mean (+) effect size (Cohen’s d) Mean (−) effect size (Cohen’s d)
Focused attention 6 0.60 ± 0.05 −0.85 ± 0.09

Mantra recitation 2 1.48 −1.11
Open monitoring 8 0.68 ± 0.12 −0.69 ± 0.14
Loving-kindness/compassion 5 0.44 ± 0.01 −0.68
All practices combined 21 M = 0.59 ± 0.02 M = −0.74 ± 0.09
Mean effect sizes ±95% confidence intervals. Note that the ‘mean’ effect sizes for mantra recitation are based on only a single study each, as are negative effect sizes for
loving-kindness/compassion meditations (cf. Table 7). Accordingly, no confidence intervals are provided since there was no variance in the estimates. Note that a total of 21
‘experiments’ were analyzed, from 17 independent studies of the 25 studies originally included in the neuroimaging meta-analysis. The remaining studies did not provide
statistical data that allowed for calculation of Cohen’s d.
(ii) Second, we discuss the results of our effect size meta-analysis Directly in line with others’ predictions (Lutz et al., 2008b), we
(Section 5.9), and its implications for the ‘practical significance’ of found activations in the premotor cortex extending into the pos-
the functional neural effects associated with meditation. We also terior dorsolateral prefrontal cortex, as well as in the dorsal
discuss the possibility of publication bias. anterior cingulate cortex. We also observed a slightly sub-threshold
(iii) Finally, several Sections (5.10–5.13) outline the limitations (88 m3 ) cluster in the dorsolateral prefrontal cortex (BA 8/9). These
of our meta-analysis, the relevance of these results to clinical con- regions are frequently activated in studies of cognitive control
ditions, and the implications for future avenues of research. that require monitoring performance, and the voluntary regula-
tion of attention and behavior, such that task-relevant actions
are selected (Carter et al., 1998; Vincent et al., 2008; Dixon and
5.2. Functional neuroanatomy of focused attention meditation Christoff, 2012; Dixon et al., 2014a). The recruitment of such
executive brain areas is consistent with a contemplative practice
5.2.1. Activations consisting largely of effortful, sustained attention with a range of
Focused attention meditation was associated with activations
in regions associated with cognitive control and self-reflection.
regulation demands. For more details, see the Supplementary dis- One of the reasons mantra is employed is to heighten the effec-
cussion. tiveness of focused attention meditation. The implication is that
by focusing upon a single salient percept (the mantra), attention
5.2.2. Deactivations is more easily sustained and distractors more easily ignored than
Consistent deactivations were observed in two major hubs of without the mantra (Goenka, 2000). The observed pattern of deacti-
the default mode network: the posterior cingulate cortex (BA vations is consistent with the idea that mantra aids in maintaining
23/31) and the posterior inferior parietal lobule (BA 39) (Buckner a sharp focus of attention, to the exclusion of incoming sensory
et al., 2008). These regions have well-established roles in mind- stimuli and awareness of one’s own body. Deactivations in audi-
wandering (Fox et al., 2015), and in particular, episodic memory tory and language-comprehension areas are also intriguing in that
retrieval, simulation of future events, and conceptual/semantic the mantra is intended as a tool for focusing attention, but is not
processing. Deactivations in these regions suggest that focused usually meant to be understood semantically.
attention meditation may diminish spontaneous thoughts regard-
ing past and future events, as well as their conceptual elaboration. 5.4. Functional neuroanatomy of open monitoring meditation
Although many regions beyond the default mode network are
implicated in mind-wandering and related spontaneous thought 5.4.1. Activations
processes, the default mode network nonetheless plays an essen- Open monitoring meditation was associated with activation
tial role (Fox et al., 2015). For more details, see the Supplementary in regions involved in the voluntary regulation of thought and
discussion. action (inferior frontal gyrus, posterior dorsolateral prefrontal
cortex/pre-motor cortex, and dorsal anterior cingulate cortex/pre-
supplementary motor area), as well as activation in the insula, a
5.3. Functional neuroanatomy of mantra recitation meditation region that plays a key role in interoceptive processing. As all of
these areas were recruited by multiple meditation techniques, they
5.3.1. Activations are discussed in more detail below in Section 5.6 on ‘Convergent
The most salient activations for mantra recitation meditation findings.’
were in the motor control network, including Broca’s area, the pre- Smaller clusters that did not exceed our threshold were
motor and supplementary motor cortices (lateral and medial BA 6), observed in left rostrolateral prefrontal cortex (BA 10) and left
as well as the putamen within the basal ganglia. Consistent with the mid-dorsolateral prefrontal cortex (BA 9/46). Together with the
goals of mantra meditation, these regions may be involved in the dorsal anterior cingulate cortex, these areas constitute key parts of
process of internally generating and staying focused on a phrase a ‘frontoparietal control’ network (Vincent et al., 2008; Spreng et al.,
within one’s mind (or recited aloud). 2010) generally involved in cognitive control, that is, coordinating
Particularly interesting was a small cluster in Broca’s area (BA and monitoring attention to both internal and external channels of
44), extending posteriorly into the face/tongue area of premotor information (Spreng, 2012; Dixon et al., 2014b). Given that open
cortex (BA 6). This area is well-known from the neuropsychologi- monitoring meditation typically involves a broad, non-judgmental
cal lesion work of Pierre Paul Broca to be critically involved in the attention to any and all mental content (including information from
motor components of speech production (Broca, 1861). Its consis- the outside world, from within the body, and also internal channels
tent recruitment during mantra recitation requires little comment. of thought and memories), it is congruous that this practice recruits
Large activations were also observed in various areas of the numerous areas from the frontoparietal control network.
basal ganglia, including bilateral clusters in both the putamen
and globus pallidus. Although these areas play a clear role in 5.4.2. Deactivations
‘reward’ (Schultz et al., 2000), they are most strongly implicated Only a single significant deactivation was observed, in the right
in highly-practiced motor movements and habit formation in gen- thalamus, and a sizeable (albeit sub-threshold) deactivation was
eral (Yin and Knowlton, 2006; Ashby et al., 2010). In a broader also observed in the left thalamus. Interestingly, in the supplemen-
sense, the basal ganglia are thought to aid in the inhibition of tal meta-analysis (which excluded practitioners with only brief
undesired movements, and in facilitating the smooth execution training in this practice), these findings were even more salient
of those movements that are desired, via tight reciprocal con- (Table S5). The thalamus is the key relay center for most incom-
nections with premotor areas (Chakravarthy et al., 2010; Stocco ing sensory information, and also functions in ‘sensory gating’
et al., 2010). These latter functions are all consistent with a medi- (i.e., the selective filtering out of certain sensory signals). Interest-
tation practice that primarily involves the deliberate execution of ingly, increased intensity of sensory gating appears to be correlated
a highly-practiced and highly-specific sequence of motor behavior with increased activity in the thalamus (LaBerge, 1990; LaBerge
(whether imagined in detail or recited aloud). For more details, see and Buchsbaum, 1990; Banich, 2004). Therefore, deactivations here
the Supplementary discussion. suggest decreased sensory gating, an interpretation that aligns well
with the goal of being open and receptive to sensory stimuli. For
5.3.2. Deactivations more details, see the Supplementary discussion.
An interesting pattern of deactivations was observed, suggesting
decreased processing of external sensory (particularly somatosen- 5.5. Functional neuroanatomy of loving-kindness and
sory) inputs. Deactivations were observed bilaterally in the anterior compassion meditation
insula, and at a sub-threshold level in the thalamus, left primary
somatosensory cortex (postcentral gyrus; BA 2), and left inferior 5.5.1. Activations
parietal lobule (BA 40). All of these areas are centrally involved Loving-kindness and compassion meditation involve cultivating
in processing and awareness of somatosensory and interoceptive positive emotions, such as joy and compassion, as well as proso-
information (Penfield and Boldrey, 1937; Bonda et al., 1995; Craig, cial skills and tendencies, such as empathic concern and altruistic
2004, 2009). Deactivations were also observed in primary auditory behavior. They focus on active ‘mentalizing’ in the form of taking
cortex (BA 41) and in the posterior area of the superior temporal the perspective of others and imagining their emotional experi-
gyrus (BA 22), both well-known to be involved in the processing and ences. Mounting evidence from behavioral studies suggests that
comprehension of sounds and language (Galaburda and Sanides, these practices can be effective, and that even brief training in
1980; Demonet et al., 1992). compassion meditation can increase empathy for, and prosocial
behavior toward, others (Leiberg et al., 2011; Condon et al., 2013; its involvement in interoception (i.e., monitoring and conscious
Kang et al., 2013; Weng et al., 2013; Lim et al., 2015). One study even awareness of internal body states, such as respiration or heart rate)
found that compassion meditation-driven changes in brain acti- (Craig, 2004, 2009; Critchley et al., 2004; Farb et al., 2013) as well
vation specifically predicted subsequent compassionate behavior as empathy (Lamm et al., 2011) and possibly even metacognition
toward an unknown victim (Weng et al., 2013). (Fleming and Dolan, 2012). Given that virtually all forms of medi-
Only three significant meta-analytic clusters were observed, tation investigated here involve some degree of monitoring of the
probably due to the low power of this analysis, which involved the body, some awareness of respiration, and basic metacognitive mon-
fewest studies (N = 5 experiments; see Table 1). One large cluster itoring, universal recruitment of insular cortex is not unreasonable.
was observed in the right somatosensory cortices (BA 2), extend- Insular cortex is not functionally homogenous, however: there
ing into the inferior parietal lobule (BA 40). A similarly located is evidence that the mid and posterior insula serve as ‘primary’
(albeit statistically non-significant) cluster was observed on the interoceptive cortex, receiving and processing direct interocep-
postcentral gyrus of the left hemisphere, centered on BA 43 and tive inputs from the thalamus (Flynn, 1999; Hua et al., 2005; Frot
extending into BA 3. These parts of parietal lobe play a well-known et al., 2007), whereas more anterior parts of the insula are prefer-
role in somatosensory processing and in creating a unified sense entially involved in conscious awareness of interoceptive signals
of the body (Penfield and Boldrey, 1937; Penfield and Rasmussen, and the integration of this information with one’s higher-order
1950), but both regions have also been implicated in empathy in goals and emotional states (Craig, 2004, 2009; Critchley et al., 2004;
a recent meta-analysis (Lamm et al., 2011) and comprehensive Critchley, 2005). In this light it is interesting that anterior insula
review (Bernhardt and Singer, 2012). Both areas are also consis- was recruited in all techniques except mantra recitation—which is
tently implicated in the perception of pain (Grant et al., 2011; Grant, the technique least focused on conscious awareness of the body,
2014), but this function may overlap with their role in empathy, and most geared toward detaching from immediate sensory stim-
as part of the proposed role of these areas in empathy may be in uli. This interpretation is speculative, of course; for more details,
specifically empathizing with the pain of others (Lamm et al., 2011). see the Supplementary discussion.
Another large cluster centered on the right anterior insula,
which has previously been implicated in quantitative meta-
analyses (Lamm et al., 2011) of the neural correlates of empathy 5.6.2. Premotor cortex and supplementary motor area
and theory-of-mind (i.e., mentalizing). Because insula activation (lateral/medial BA 6)
was observed across multiple meditation categories, its possible Three categories of meditation (the exception being loving-
role(s) in loving-kindness meditation is discussed further in the kindness/compassion) showed large meta-analytic clusters in
section on convergent findings (Section 5.6). For more details, see posterior dorsolateral prefrontal cortex/premotor and supplemen-
the Supplementary discussion. tary motor cortices (BA 6). Although these areas are traditionally
considered ‘motor’ regions, a wealth of recent work has demon-
5.5.2. Deactivations strated that they also play a role in many cognitive functions
No significant deactivations were observed; for more details, see (e.g., Hanakawa et al., 2002; Tanaka et al., 2005). Specifically, it
the Supplementary discussion has been proposed that these areas could play a role analogous
to their motor function of manipulating objects in the physical
5.6. Convergent findings across meditation categories world—namely, the mental manipulation of psychological content
and memory (Hanakawa et al., 2002). Functional neuroimaging
Relatively few regions showed significant clusters across mul- studies have shown activation in these areas during a variety of
tiple meditation categories, supporting the central hypothesis of complex mental tasks, including working memory (Fiez et al., 1996;
this paper that distinct meditation styles recruit largely non- Owen and Evans, 1996), attentional control (Hopfinger et al., 2000;
overlapping neural networks. Only a single area (insular cortex) Boussaoud, 2001), mental imagery (Mellet et al., 1996), and concep-
was recruited by all four meditation categories. Some additional tual reasoning (Rao et al., 1997). Given the apparent role of these
consistencies across meditation types were observed, however; regions in diverse higher-order cognitive functions and attention
each area is considered in turn below. regulation—the core of all meditation techniques—their recruit-
ment across numerous styles of meditation appears reasonable.
5.6.1. Insular cortex Speculatively, then, these regions may interact with other brain
Meta-analytic activations were observed in insular cortex in all areas more specifically tied to particular meditation techniques,
four meditation categories explored here. For focused attention, assisting in the intentional regulation of attention and other cogni-
a small (non-significant) cluster was observed in the left anterior tive processes specific to the practice at hand.
insula, as well as a somewhat larger (also non-significant) cluster
in the left mid-insula. For mantra recitation, a very small (non-
significant) cluster was observed in the left mid-insula/claustrum. 5.6.3. Dorsal anterior/mid cingulate cortex
For open monitoring meditation, a large (significant) cluster was In focused attention and open monitoring meditation, we
observed in the left anterior/mid-insula, as well as two smaller observed significant activation clusters that bordered both the sup-
(non-significant) clusters in the right mid- and posterior insula. plementary motor areas (discussed in the previous section) and
For loving-kindness meditation, a large (significant) cluster was the dorsal anterior cingulate cortex. Congruent with these find-
observed in the right anterior insula. The insula therefore appears to ings, consistent structural alterations in dorsal anterior cingulate
be reliably recruited in two categories of meditation we examined, cortex were observed in our recent meta-analysis of morphomet-
and shows evidence for recruitment in the two other styles. ric neuroimaging of meditation (Fox et al., 2014). Despite numerous
These findings suggest an important role for the insula across functional roles, the dorsal anterior cingulate is clearly a key player
numerous meditative practices. They are also consistent with our in the brain’s ability to regulate attention and emotion, as well as
recent meta-analysis of morphometric neuroimaging of medita- monitor performance (Bush et al., 2000; Posner et al., 2007). Activa-
tion, which collapsed across many meditation styles and found that tions here across the two styles of meditation that most explicitly
differences in insula were the most consistent structural alterations involve such regulation and monitoring is therefore sensible. For
(Fox et al., 2014). Although the insula has many roles, probably further discussion, see the individual sections on focused attention
the most consistently replicated and thoroughly investigated is and open monitoring.
5.6.4. Frontopolar cortex/rostrolateral prefrontal cortex when not engaging in meditation, which led to recruitment of all
We observed a fairly large but sub-threshold cluster of the regions commonly implicated in pain processing, during med-
activation in the left frontopolar cortex (BA 10) for open moni- itation there was a marked deactivation of these classic pain areas,
toring meditation. For loving-kindness/compassion meditation, we including the thalamus, secondary somatosensory cortex, insula,
observed a small (non-significant) cluster in the superior portion of and cingulate cortex. Again, none of the four practice types meta-
left frontopolar cortex (BA 10), extending somewhat into the dor- analyzed here bears much resemblance to this pattern (and open
solateral prefrontal cortex (BA 9). Although neither of these results monitoring seems almost antithetical to it—in line with the meth-
exceeded our stringent significance thresholds, nonetheless these ods and motivations underlying each practice).
findings are in line with our recent meta-analysis of morphomet- Finally, a single study investigating non-dual awareness in highly
ric (i.e., structural-anatomical) neuroimaging studies of meditation experienced Tibetan Buddhist practitioners reported evidence that
(Fox et al., 2014), and therefore warrant a brief discussion. this practice decreased the anti-correlation between activity in
In our morphometric meta-analysis, we reported anatomical ‘extrinsic’ and ‘intrinsic’ networks of the brain, which are argued to
increases (e.g., increased cortical thickness; Lazar et al., 2005) in mediate attention to the external environment and internal states,
this area across three fairly divergent schools of practice (Fox et al., respectively (Josipovic et al., 2011). The authors created numer-
2014). This region is known to play a key role in meta-awareness ous regions-of-interest throughout the brain corresponding to an
and metacognitive capacity generally (e.g., (Fleming et al., 2010, ‘extrinsic’ network for attention to external stimuli, and an ‘intrin-
2012; McCaig et al., 2011; Fleming and Dolan, 2012; Fox and sic’ network thought to be involved in internal streams of thought
Christoff, 2015)), but it also appears to be involved in the eval- (which roughly parallels the default mode network). The extrinsic
uation of self-generated information (Christoff et al., 2003) and and intrinsic networks are thought to show anti-correlated patterns
in the processing of complex and abstract information generally of functional connectivity during either external tasks or atten-
(Christoff et al., 2001, 2009). Frontopolar cortex is also a key node tion to internal streams of thought (e.g., during mind-wandering)
of the frontoparietal control network (Vincent et al., 2008) and may (Fox et al., 2005). The authors examined the extent to which this
play an important role in switching attention between external and anti-correlated functional connectivity might be attenuated by
internal channels (Ramnani and Owen, 2004; Burgess et al., 2007). non-dual awareness meditation, which broadly aims at reducing
The specific functions of the frontopolar cortex in various forms of the sense of a division between external and internal. During non-
meditation remain an open question. dual awareness meditation, the anti-correlation between these two
networks was markedly attenuated compared to either focused
5.7. Other meditation practices: further evidence for attention meditation or passive fixation. The authors interpreted
dissociability at the neural level these results as evidence of the effectiveness of non-dual awareness
practice at dissolving the boundary between internal and exter-
In the Introduction, we noted that there are several other major nal experiences (i.e., subject-object duality; Josipovic et al., 2011).
categories of meditation wherein research remains too sparse for In line with our central hypothesis and meta-analytic results, the
inclusion in a meta-analysis (Section 1.3). These categories include authors also suggested that their results indicated that non-dual
visualization (generating and sustaining complex visual imagery), awareness practice is differentiable at the neural level from other
non-dual awareness (eroding the boundary between subject and meditations, such as focused attention and open monitoring med-
object), and pratyahara (dampening of sensory inputs). Neural cor- itation (Josipovic et al., 2011).
relates of these practices have been reported in isolated studies, Although the evidence regarding these other forms of medi-
and interestingly, provide further support for the hypothesis that tation remains tentative, it seems clear that further research is
distinct psychological () practices are likewise dissociable at the warranted into various other techniques that have not yet been
neurobiological () level (Cacioppo and Tassinary, 1990). intensively investigated. Based on our meta-analytic findings, as
For instance, a single study has examined the neural corre- well as the studies just reviewed, we suspect that future work will
lates of visualization meditation (Lou et al., 1999). In this study, discover and refine dissociable neural correlates associated with
practitioners were guided through “visual imagination of a sum- these various psychologically unique meditation practices.
mer landscape with forests, streams, and meadows with cattle”
(Lou et al., 1999). Although this particular imagined content is not 5.8. Are contemplative practices dissociable by
typical of visualization meditation practices (which tend to focus electrophysiological and neurochemical measures?
on more explicitly religious themes), nonetheless the results are
telling. Visualization meditation resulted in significant recruitment Virtually all of the meta-analytic results and discussion in this
(among other areas) of the fusiform gyrus (definitively linked to review have relied on brain signals indicative of cerebral blood
higher-order visual processing; McCarthy et al., 1997; McCandliss flow (PET) or blood oxygenation (fMRI), but there are numerous
et al., 2003), the parahippocampal gyrus (linked to visual ‘place’ other ways of characterizing brain function, most notably via elec-
perception; Epstein et al., 1999), and nearly universal recruitment trophysiological activity (via electroencephalography, for instance;
of medial and lateral occipital areas (Lou et al., 1999). Intriguingly, EEG) and at the neurochemical level (i.e., the tonal and phasic rates
primary visual cortex (V1), which is not required for the creation of release of various neurotransmitters across different areas of
of visual imagery and visual dreaming (Solms, 1997), was essen- the brain). For more details on how these possibilities have been
tially the only occipital area not strongly recruited (Lou et al., 1999). applied in meditation research, see the Supplementary discussion.
These results, although drawn from only a single study, support the
idea that practitioners were successfully engaging in such a visu- 5.9. Do functional neural effects associated with meditation have
alization practice. More importantly, for our purposes, they are practical significance?
utterly distinct from any of the broad patterns of neural activity
reported in this meta-analysis, further supporting the contention Our effect size meta-analyses found that the four primary
that differing mental practices have dissociable and identifiable categories of meditation examined were all associated with
neural correlates. approximately ‘medium’ effect sizes (mean Cohen’s d = 0.59 for acti-
A case study involving a single highly-experienced practitioner vations and −0.74 for deactivations; for details see Tables 7 and 8).
engaging in a pratyahara (sense-withdrawal) practice is similarly Although we find these results both reasonable (in magnitude) and
informative (Kakigi et al., 2005). Compared to painful stimulation encouraging, we reiterate that the ‘meaningfulness’ of effect sizes is
tions) and ‘negative’ effect sizes (brain deactivations). The simplest

interpretation of these results is that fairly serious publication bias
(i.e. non-publication of null and ‘non-significant’ findings) afflicts
this field. Even if meditation always and truly results in significant
effects on brain activation or deactivation, given that 78 functional
neuroimaging studies had been published at the time of writing,
we would expect at least a few null results to have been obtained
and reported simply by chance alone. An alternative (albeit related)
explanation is that smaller effect sizes were not observed precisely
because of the inflationary bias inherent in neuroimaging reporting
of effect sizes (see Section 2.5). Although this possibility presents
at least as large a problem as does publication bias, it shifts the
emphasis from the limitations of the current scientific publication
model (which places undue emphasis on ‘positive’ findings) to the
limitations of neuroimaging data analysis and reporting practices.
In either case, the lack of reported small or null effects in this liter-
ature (even after our severe deflationary procedures) suggests bias
at some stage of study design, analysis, or reporting.
Fig. 6. Funnel plot showing weighted means of both negative and positive effect That said, for both negative and positive effect sizes, the
sizes. The two anchor lines represent the means for all (−) and (+) effect sizes
expected funnel plot pattern was generally observed: that is, with
(−0.74 and +0.59, respectively) weighted by sample size. Positive effect sizes relate
to brain activations during meditation, whereas negative effect sizes represent brain increasing sample size, the variability among effect sizes reported
deactivations. See Tables 7 and 8 for details. decreased drastically; and notably, the most extreme effect sizes
were always reported in studies with the smallest sample sizes. This
suggests, encouragingly, that despite potential publication bias or
ultimately arbitrary even where their use is highly developed (e.g., inflationary bias due to neuroimaging analysis methods, nonethe-
in the behavioral, social, and medical sciences; Cohen, 1992). This less studies with larger samples tend to converge on similar and
is all the more true in neuroimaging, where their calculation and more reasonable (medium) effect sizes. Although such a conclu-
interpretation are poorly understood (see Supplementary meth- sion is tentative, the results to date (Fig. 6) suggest that a sample
ods). Nonetheless, some general observations can be made. size of approximately n = 25 is sufficient to reliably produce effect
First, positive and negative effect sizes (representing activa- sizes that accord with those reported in studies with much larger
tions and deactivations associated with meditation, respectively) samples (up to n = 46).
are similar in magnitude (Tables 7 and 8; Fig. 6). This is in line
with our meta-analytic neuroimaging findings (where brain deac- 5.10. Meditation, the brain, and clinical disorders
tivations were nearly as common and significant as activations) and
supports the idea that the successful execution of a particular med- Although the central aim of this meta-analysis was to better
itation may depend as much on which brain areas are disengaged delineate the neural correlates of meditation in healthy practition-
as on which are actively recruited. ers, the potential effects of meditation on various physical and
Second, the approximately ‘medium’ magnitude of the effects psychological clinical conditions is of enormous interest (Gotink
suggests that the functional neural effects of meditation result in et al., 2015) and therefore warrants a brief discussion. To our
roughly a one-half standard deviation increase or decrease in brain knowledge, only five studies to date (Table S1) have examined the
activity, relative to various other baseline conditions, or novices relationship between meditation, brain activity, and clinical disor-
attempting to execute the same practice as experts. ders of some kind. One study investigated social anxiety disorder
Third, differing meditation categories showed largely similar (Goldin and Gross, 2010) and another bipolar disorder (Ives-
mean effect sizes—roughly one half to two-thirds of a standard Deliperi et al., 2013); one examined patients with breast cancer
deviation (Cohen’s d ∼0.50–0.67; see Table 8). The exception was (Monti et al., 2012); one involved chronic tobacco smokers dur-
mantra recitation, which appeared to result in larger effects for ing a period of abstinence from smoking (Westbrook et al., 2013);
both activations and deactivations (Cohen’s d > 1.0). This apparently and one recruited patients with mild cognitive impairment (Wells
large difference is highly questionable, however: first, these ‘mean’ et al., 2013). The heterogeneity of these investigations precludes
(+) and (−) effect sizes across studies are in fact simply mean effect any quantitative meta-analytic procedure, but nonetheless these
sizes for a single study each, as only two studies contributed to the preliminary explorations are a very important step toward exam-
effect size meta-analysis for mantra recitation (see Tables 7 and 8). ining the potential of meditation to ameliorate such conditions,
Moreover, the two studies that contributed (Davanger et al., 2010; as well as understanding the cognitive and neural mechanisms by
Kalyani et al., 2011) have among the smallest sample sizes of any which these improvements might take place. Considering the fairly
studies we investigated here (see Table 7). This makes these reports ample evidence from psychological research that meditation can
highly susceptible to the inflationary biasing of effect sizes we have salutary effects on affective characteristics (Sedlmeier et al.,
discussed at length in the Supplementary methods. We therefore 2012; Goyal et al., 2014), further investigations into its potential
conclude that it would be highly premature to argue that mantra benefits for clinical conditions, and the brain mechanisms related
meditation results in larger effects than other forms of medita- to these benefits, are of paramount importance.
tion practice. A more plausible interpretation is that inflationary
bias was larger in these studies, and the true effect size of mantra 5.11. Meta-analytic limitations
recitation is in fact comparable to the other practices examined
here. First, it should be reiterated that the four categories employed in
Fourth, a funnel plot (scatterplot of effect size vs. sample size; our synthesis combined meditation practices which, although sim-
Fig. 6) showed a conspicuous dearth of null results or small effect ilar, are not necessarily identical. Moreover, there are several other
sizes (even after our multiple deflationary adjustments of originally major categories of meditation that could not be quantitatively
reported effect sizes), for both ‘positive’ effect sizes (brain activa- examined here (e.g., non-dual awareness and sensory-withdrawal
Fig. 7. Summary of meta-analytic activations and deactivations for four major categories of meditation.
Activations are represented by filled circles, deactivations by circles containing downward arrows. Regional labels are approximate, and are shown for illustrative purposes
only. For details, see tables of results, main figures, and supplementary figures. ACC: anterior cingulate cortex; aIns: anterior insula; BG: basal ganglia; Broca: Broca’s area;
dACC: dorsal anterior cingulate cortex; DLPFC: dorsolateral prefrontal cortex; IFG: inferior frontal gyrus; IPL: inferior parietal lobule; mIns: mid-insula; PCC: posterior cingulate
cortex; PMC: premotor cortex; RLPFC: rostrolateral prefrontal cortex; RMPFC: rostromedial prefrontal cortex; SII: secondary somatosensory cortices; SMA: supplementary
motor area.
practices), due to the paucity of empirical data to date (see Section the same practitioners and/or within the same study, and using
1.3). more homogenous baseline/control conditions (e.g., Brewer et al.,
Second, despite inclusion of all eligible studies to date, the 2011). Such an approach would control for many of the poten-
present analysis relies on relatively few research reports (n = 25; tial confounds that plague comparison across different studies,
see Table 1). We also combined results from long-term practi- conducted by different researchers, investigating different sam-
tioners with those of novices undergoing short-term training in ples of practitioners. Taken together, the present results should be
an effort to include all available data (see Table 2). This may be interpreted with caution: a clearer understanding of the major cat-
problematic because, as noted earlier, the diverse goals of medita- egories of meditation, as well as their subtler subdivisions, awaits
tion practices can take considerable time to be realized. It might further research undertaken with more directly comparable meth-
therefore be expected that the distinctions between brain activa- ods, baselines, and populations.
tion patterns driven by particular forms of meditation may be even
sharper among expert vs. novice practitioners, or that brain activity 5.12. Other methodological challenges in the neuroimaging of
in novices might otherwise influence or drive the effects observed meditation
here. To mitigate these concerns, however, we conducted a series
of four supplemental meta-analyses where we excluded studies of Other major concerns have less to do with meta-analytics or
short-term meditation training, and found little evidence that this statistical issues, but rather relate specifically to the investigation
was the case. Meta-analytic results for each meditation category of meditation practitioners. Several important themes are outlined
employing only long-term practitioners were negligibly different here.
from our main meta-analyses, except in the case of open moni-
toring meditation. In the case of open monitoring meditation, the
5.12.1. Lifestyle variables
general pattern of results was also similar, but the precise locations
The first concern is the general and unknown effect of lifestyle.
and significance levels of various clusters were different (for details,
Highly-experienced meditation practitioners are likely to have
compare Table 5 with Table S4).
more meager (calorie-restricted) and more vegetarian diets; are
Third, study methods and practitioner samples were highly
less likely to have children or long-term regular employment; and
heterogeneous. Both meditation and baseline conditions differed
probably sleep less than the average person (Britton et al., 2014).
considerably across studies (even within a single meditation cat-
These are just a few major examples; an array of other factors might
egory), as did meditation experience level (even among expert
be relevant. Different meditation traditions emphasize such factors
practitioners).
to varying degrees, and it is certainly conceivable that major differ-
Fourth, numerous and serious difficulties attend the calculation
ences in diet, stress, and sleeping patterns could have effects on
and interpretation of effect sizes in neuroimaging data. Our results
brain structure and function. Control subjects in the studies meta-
in this sphere, and their bearing on the question of the ‘practi-
analyzed here, however, typically only control for simple variables
cal significance’ of meditation’s effects on brain function, need to
like age, sex, and sometimes handedness, and are therefore unable
be viewed in light of the many caveats and limitations we have
to account for these lifestyle variables. Until these and other fac-
discussed at length in the Supplementary methods.
tors are carefully controlled for, the potentially confounding effects
Future work may be better able to mitigate many of these limi-
of non-meditation-specific aspects of the contemplative lifestyle
tations by directly examining multiple forms of meditation within
cannot be ruled out or fully understood (Fox et al., 2014).
5.12.2. Pre-existing differences in brain structure and function instance measuring changes in functional connectivity at baseline
A second general concern is the possibility of pre-existing dif- (in the absence of any task or stimuli).
ferences in brain structure or function in advanced meditation
practitioners (Fox et al., 2014). Given the rigorous and peculiar 5.12.5. Integration of functional neuroimaging methods with
demands of spending thousands of hours in solitary, minimally- behavioral measures
stimulating environments and unusual (for Western practitioners) As we discussed in our previous meta-analysis of morphometric
and often painful physical postures, it seems highly likely that neuroimaging studies of meditation (Fox et al., 2014), alterations in
long-term contemplative practice may preferentially attract per- brain structure and function are of little value (beyond basic scien-
sons with a proclivity toward solitude and sustained attention to tific interest) if they are not accompanied by changes in behavior
inner mental life. Support for this hypothesis comes from a recent or cognitive-emotional experience and wellbeing (Fox et al., 2016).
study that was able to predict subsequent practice of two different We discuss this issue and some of the findings we reviewed related
forms of meditation (mindfulness and compassion) from baseline to brain-behavior correlations in the Supplementary discussion.
brain activity alone (Mascaro et al., 2013). Cross-sectional studies
(which form a large part of this meta-analytic dataset) cannot con- 5.13. Summary and conclusions
trol for such pre-existing differences. On the other hand, studies
involving short-term training of novices in pre-post designs cannot The present meta-analysis is the first to report dissociable
examine the potentially subtle effects that long-term and commit- patterns of brain activation and deactivation underlying four
ted practice may have on brain function. Future work will have major categories of meditation (focused attention, mantra recita-
to strike a balance somewhere between the limitations of these tion, open monitoring, and loving-kindness/compassion practices),
two general approaches, and further studies of pre-existing (and and provides suggestive evidence for the dissociability of three
potentially predictive) differences in brain structure and function additional meditation types (visualization, sense-withdrawal, and
in meditation practitioners would be highly beneficial (cf. Mascaro non-dual awareness practices). Our synthesis provides preliminary
et al., 2013). evidence for largely dissociable neural substrates which, broadly
speaking, are consistent with the methods, aims, and putative psy-
chological results of the practices examined. Collectively, these
5.12.3. ‘Overshadowing’ results suggest that while several regions may be equally involved
A third and major concern is what we call ‘overshadowing’—the in many forms of contemplative practice (Sperduti et al., 2012),
possibility that a practitioner’s background experience with a vari- differences in neural activity greatly outnumber similarities. Com-
ety of meditation techniques might color the neurophysiological monality across meditation categories is the exception rather than the
results obtained during the brief ‘state’ investigations of isolated rule. Importantly, deactivations almost always accompany activa-
meditation techniques that are typical of the experiments exam- tions (though they are usually fewer in number, less substantial in
ined here. This issue is discussed further in the Supplementary terms of cluster size, and less consistent across studies in terms of
discussion. peak likelihood value). This suggests that for the effective execu-
tion of various meditation practices, disengaging particular brain
regions may be nearly as important as engaging others.
5.12.4. Enduring trait differences in brain and cognitive function After careful corrections and adjustments for suspected infla-
Closely related to the overshadowing effect are unknown trait tionary biases, effect size meta-analysis suggests that meditation
differences. By ‘overshadowing’ we mean the possibility that accu- practices tend to yield approximately medium-sized effects. These
mulated trait differences might overshadow (or interact with) results suggest that meditation’s relationship with brain function
brain activations and deactivations directly associated with a may be practically significant, but there appears to be a long way
brief, induced state of meditation in neuroimaging investiga- to go in the statistical theory of neuroimaging before these find-
tions. But although these putative trait differences are a potential ings can be interpreted with confidence. Moreover, it is imperative
confounding factor for our purposes here (investigating medi- that neuroimaging investigations of meditation begin to rigorously
tation state effects), enduring trait changes are an explicit goal relate findings to behavioral and self-reported outcomes, in order to
of most meditation traditions, and are a worthy object of study understand how (and to what extent) these changes in brain activ-
in their own right. Here, we chose to exclude such studies ity are related to actual cognitive-affective change and benefits in
from our meta-analysis (criterion iv; see Section 2.1.2), mainly practitioners.
because of the enormous methodological challenges involved in Many have understandably viewed the nascent neuroscience of
synthesizing across so many disparate tasks and contemplative meditation with skepticism (Andresen, 2000; Horgan, 2004), but
traditions. Although our meta-analysis was also affected by dif- recent years have seen an increasing number of high-quality, con-
fering baseline/comparison conditions, and potentially also an trolled studies that are suitable for inclusion in meta-analyses and
‘overshadowing’ effect, nonetheless the meditation states inves- that can advance our cumulative knowledge of the neural basis of
tigated were constant and consistent for each analysis (focused various meditation practices (Tang et al., 2015). With nearly a hun-
attention, mantra meditation, etc.). The major challenge facing dred functional neuroimaging studies of meditation now reported,
any effort to synthesize trait effects associated with meditation we can conclude with some confidence that different practices
practice is that even this limited homogeneity (constant medita- show relatively distinct patterns of brain activity, and that the
tion states, despite differing samples and comparison conditions) magnitude of associated effects on brain function may have some
is absent. Not only is there great heterogeneity among the tasks practical significance. The only totally incontrovertible conclusion,
on which meditation-related traits are being investigated (e.g., however, is that much work remains to be done to confirm and
emotion regulation, perceptual discrimination, etc.), no consistent build upon these initial findings.
meditation state is being induced, and the practitioners employed
will invariably possess heterogeneous contemplative backgrounds Acknowledgments
with respect to the proportion, and absolute amount, of time spent
in various different practices. One way to begin to examine such We thank Dr. Antoine Lutz and Dr. David M. Fresco for very help-
trait differences, which sidesteps these problems, would be to com- ful discussions of the methods and results of this meta-analysis, and
pile data on long-term practitioners during the ‘resting’ state, for Arianne Herrera-Bennett for assistance in creating Fig. 7. We also
thank Dr. Norman Farb for providing further details from previously Craig, A., 2004. Human feelings: why are some more aware than others? Trends
unpublished data that contributed to the effect size meta-analysis. Cogn. Sci. 8, 239–241.
Craig, A., 2009. How do you feel—now? The anterior insula and human awareness.
This work was supported in part by Natural Sciences and Engineer- Nat. Rev. Neurosci. 10.
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awarded to K.C.R.F. and M.L.; NSERC Canada Graduate Scholarships supporting interoceptive awareness. Nat. Neurosci. 7, 189–195.
Critchley, H.D., 2005. Neural mechanisms of autonomic, affective, and cognitive
awarded to M.L.D. and M.E.; a University of British Columbia Affili- integration. J. Comp. Neurol. 493, 154–166.
ated Doctoral Fellowship awarded to J.L.F.; and NSERC and Canadian Cumming, G., 2012. Understanding the New Statistics: Effect Sizes, Confidence
Institute of Health Research (CIHR) grants awarded to K.C. Intervals, and Meta-analysis. Routledge.
Cumming, G., 2013. The new statistics why and how. Psychol. Sci., http://dx.doi.
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Appendix A. Supplementary data
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Neuroscience and Biobehavioral Reviews 65 (2016) 208–228

Contents lists available at ScienceDirect

Neuroscience and Biobehavioral Reviews

Functional neuroanatomy of meditation: A review and meta-analysis

2. Meta-analytic methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213

1. Introduction and mounting empirical evidence suggests that meditation indeed

synthesis, we sought to build on prior reviews and meta-analyses

Table 1 late to clinical signiﬁcance or relevance (Rosnow and Rosenthal,

Study Practice N (M/C) Tradition Expertise Average Experience

Lou et al. (1999) OM + LK 9/– Yoga Nidra LTP >5 years

All peak voxel coordinates are reported in Montreal Neurolog-

Region Cluster Size Side Peak Coordinates Peak ALE value

Brefczynski-Lewis et al. (2007) 30 FA 0.51 –

Focused attention 6 0.60 ± 0.05 −0.85 ± 0.09

All practices combined 21 M = 0.59 ± 0.02 M = −0.74 ± 0.09

tions) and ‘negative’ effect sizes (brain deactivations). The simplest

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