**9. Conclusions**

Compared to previous neuroimaging techniques, fMRI technology has generally improved our understanding of neural networks involved in mental imagery in all sensory modalities. The higher sensitivity of fMRI allowed researchers to better differentiate between primary and secondary sensory cortices, whereas the extreme flexibility of scanning procedures allowed the implementation of different imagery paradigms, which likewise has implications for the imagery debate issue.

Several, several conclusions can be made from the fMRI literature reviewed above. First, the involvement of the primary sensory cortices varies according to the imagery modality investigated, and may be modulated by different factors, such as the individual differences in generating mental images or expertise, tasks, types of paradigms, or analyses employed. Second, the activation of the secondary sensory cortices in all imagery modalities is more consistent, as well as the recruitment of pre-frontal areas or associative cortices, the former plausibly participating in the top-down processes which are executed for retrieving information from long-term memory, the latter facilitating supra-modal processing of sensory information. This picture is partially in contrast with the perceptual approach, which assumes that mental images correspond with percepts and events also in terms of both mechanisms and processes used. It also supports the propositional approach, which assumes an amodal format for all types of mental images. Indeed, when generating mental images, the human brain seems to rely mostly on secondary sensory cortices rather than primary ones, clearly indicating that imagery and perception rely on overlapping but dissociable neural networks.

Looking separately at the relevant fMRI literature on imagery modalities, more detailed information can be obtained. First, fMRI showed that under specific conditions, visual imagery corresponds to the isolated activation of visual cortical areas, including the primary one, with concurrent deactivation of irrelevant sensory processing (Amedi et al., 2005). However, the overlap between visual imagery and visual perception is more consistent in the frontal and temporal cortices (Ganis et al., 2004) or ventro-temporal cortex (Reddy et al., 2010), rather than in the occipital cortex. The insights for visual imagery cannot be simply generalized, given the different sensory characteristics of percepts, as well as the roles that each imagery system can play in the cognitive system.

Indeed, fMRI literature showed that auditory verbal imagery relies mostly on the secondary auditory cortex, left inferior frontal cortex, supplementary motor area, and inferior parietal areas, rather than the primary auditory cortex. The former regions were found to be involved in the verbal perception domain: particularly, left pre-frontal areas mediate phonological recoding (Thierry et al., 1999), whereas the supplementary motor area is important for silent articulation or inner speech, and posterior superior temporal gyrus or temporo-parietal junction are involved in the acoustic-phonetic feature-based processing (Scott & Johnsrude, 2003).

Moreover, auditory imagery of environmental sounds never yielded activation in the primary auditory cortex. Even when the sparse temporal sampling technique was used to

sensations of hungry, thirsty, cold, drunkenness, etc. Very few fMRI studies were devoted to proprioceptive imagery. Olivetti Belardinelli et al. (2004a) did not show any activation of the primary somatosensory cortex during proprioceptive imagery, whereas Olivetti Belardinelli et al. (2009) revealed that high vivid participants activated the right post-central gyrus (BA

Compared to previous neuroimaging techniques, fMRI technology has generally improved our understanding of neural networks involved in mental imagery in all sensory modalities. The higher sensitivity of fMRI allowed researchers to better differentiate between primary and secondary sensory cortices, whereas the extreme flexibility of scanning procedures allowed the implementation of different imagery paradigms, which likewise has

Several, several conclusions can be made from the fMRI literature reviewed above. First, the involvement of the primary sensory cortices varies according to the imagery modality investigated, and may be modulated by different factors, such as the individual differences in generating mental images or expertise, tasks, types of paradigms, or analyses employed. Second, the activation of the secondary sensory cortices in all imagery modalities is more consistent, as well as the recruitment of pre-frontal areas or associative cortices, the former plausibly participating in the top-down processes which are executed for retrieving information from long-term memory, the latter facilitating supra-modal processing of sensory information. This picture is partially in contrast with the perceptual approach, which assumes that mental images correspond with percepts and events also in terms of both mechanisms and processes used. It also supports the propositional approach, which assumes an amodal format for all types of mental images. Indeed, when generating mental images, the human brain seems to rely mostly on secondary sensory cortices rather than primary ones, clearly indicating that imagery and perception rely on overlapping but

Looking separately at the relevant fMRI literature on imagery modalities, more detailed information can be obtained. First, fMRI showed that under specific conditions, visual imagery corresponds to the isolated activation of visual cortical areas, including the primary one, with concurrent deactivation of irrelevant sensory processing (Amedi et al., 2005). However, the overlap between visual imagery and visual perception is more consistent in the frontal and temporal cortices (Ganis et al., 2004) or ventro-temporal cortex (Reddy et al., 2010), rather than in the occipital cortex. The insights for visual imagery cannot be simply generalized, given the different sensory characteristics of percepts, as well as the roles that

Indeed, fMRI literature showed that auditory verbal imagery relies mostly on the secondary auditory cortex, left inferior frontal cortex, supplementary motor area, and inferior parietal areas, rather than the primary auditory cortex. The former regions were found to be involved in the verbal perception domain: particularly, left pre-frontal areas mediate phonological recoding (Thierry et al., 1999), whereas the supplementary motor area is important for silent articulation or inner speech, and posterior superior temporal gyrus or temporo-parietal junction are involved in the acoustic-phonetic feature-based processing

Moreover, auditory imagery of environmental sounds never yielded activation in the primary auditory cortex. Even when the sparse temporal sampling technique was used to

2/3) during proprioceptive imagery in comparison with low-vivid participants.

**9. Conclusions** 

implications for the imagery debate issue.

dissociable neural networks.

(Scott & Johnsrude, 2003).

each imagery system can play in the cognitive system.

ensure that the acoustic scanner noise did not lead to interferences with the auditory perception and imagery conditions, the primary auditory cortex was not activated (Bunzeck et al., 2005). This pattern of results shows that when imaging environmental sounds the topdown process is initiated mostly by the inferior frontal gyrus and insular regions, and manages to activate only secondary auditory areas.

Still, the fMRI literature on musical imagery, with the exception of Kraemer et al. (2005), showed no activation of the primary auditory cortex, even when enrolling musicians. Given the pattern of activation yielded by musical imagery, an associative network independent of primary sensory-motor and auditory activity is likely representing the cortical elements most intimately linked to music production. In addition, considering that both amateurs and professional musicians did not report any activation in the primary auditory cortex, and that motor areas were not recruited, either, during musical imagery involving mental performances, it is highly probable that these areas become tightly coupled with executed activities during musical training. Finally, given that musical imagery in professional musicians revealed more anterior cerebellar activations, it is possible to conclude that musical imagery involves the recruitment of stored movement programs of sequential finger movements (Lotze et al., 2003).

The extent to which the primary motor cortex is involved in motor imagery remains unclear. However, recent fMRI studies have highlighted the importance of the supplementary motor area, which seems to be involved in suppressing the activity of the primary motor cortex, and consequently, movements that are not intended to be performed. This would mean that motor imagery and motor execution do not rely on the same neural network.

Regarding olfactory imagery, fMRI results are in line with behavioral findings: if images of smells are cued by verbal labels it is hard to detect the activation of the primary olfactory cortex. On the contrary, if odors are perceptually encoded, it is possible to find activation in the piriform cortex when generating images of smells. However, the activity in the olfactory primary cortex was found to be modulated by the hedonic pattern of stimuli to be imagined, and by expertise in generating or using olfactory images. This means that olfactory imagery may activate olfactory areas under specific conditions, basically when people really evoke images of smells.

The literature on gustatory, tactile, and proprioceptive imagery is more consistent. All three of these modalities seem to rely on the primary sensory cortices. Nevertheless, given the scarcity of studies carried out on these imagery systems, especially on the proprioceptive imagery, any conclusion would be premature.

In conclusion, fMRI technique has helped to clarify the neural circuits of mental imagery, but a lot a work needs to be still done. For example, the role of certain cortical areas remains unclear, as does the connectivity among cortices in mental imagery. Research is trying to clarify this issue in respect to the motor imagery modality, but it should also be addressed in respect to the others. It is also important to explore the common cortical networks involved in all imagery modalities to clarify the extent to which mental imagery relies on non-sensory cortices. Indirectly, this information would contribute to improve our understanding of the imagery debate, as having access to the common areas shared by all imagery modalities provides an understanding of the extent to which imagery involves propositional encoding. Currently, only Olivetti Belardinelli et al. (2001, 2004b) have attempted to investigate this issue. Though there has been some difference in terms of anatomical extension, results revealed that the generation of images in all sensory modalities activated the left inferior temporal cortex (including the left fusiform gyrus) and the bilateral inferior parietal lobule.

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**Part 5** 

**Post Traumatic Stress Disorder** 

