In 1995-96, when Andreas Tolias arrived at MIT to work in the laboratory of Peter Schiller, it was clear that he would become an exceptional neuroscientist. One of the first recommendations Andreas made was to have me read up on the neurophysiology of time perception. Having done his undergraduate at Cambridge University, he overlapped with the renowned physicist, Stephen Hawking, who had much to say about time and space. As it was described to me, Hawking had a propensity of driving his motorized wheelchair aggressively such that those walking about would be made to jump aside as he charged passed. Great men/women often have idiosyncrasies, but if they are not given unlimited power the world is able to tolerate them. Albert Einstein understood his idiosyncrasies for he never became the president of Israel—albeit some have suggested his decision was based on his morals.

So, what about timing and the brain? In mammals, neocortical oscillations, especially in regions associated with olfaction, are synchronized to the rate of breathing (Adrian 1942, 1950). When human subjects breathe through the nose the effect of respiration on cortical oscillations (~ 0.16 to 0.33 Hz/period of 6 to 3 seconds, respectively) measured subdurally from the orbital cortex (including the periform cortex) and the temporal lobes (including the hippocampus and amygdala) is synchronized with inhalation (Zelano, Gottfried et al. 2016). This synchronization is phase-locked with cortical oscillations of higher frequency such as theta and gamma whereby breathing rate can act as a neural clock (Fontanini and Bower, 2006; Moore et al., 2013; Ito et al., 2014). However, if breathing is now made to occur through the mouth then the relationship between breathing and neural oscillations disappears (Zelano, Gottfried et al. 2016). Thus, during periods of focused exertion, such as running or swimming, the neocortical rhythms are decoupled from respiration and this mediation is subsumed by subcortical networks that are under automatic control (e.g., the cerebellum, Tehovnik, Hasanbegović, Chen 2024) (see Footnote 1).

This neocortical decoupling would occur as athletes like Usain Bolt traverse a 100-meter track such that two inhalations are sufficient to complete one track-length (i.e., two breaths per 42 strides, Usain Bolt, Wikipedia 2024). Information about stride is conveyed to the cerebellum via the proprioceptive, mossy fibre system (Fuchs and Kornhuber 1969; Gibson et al. 2004) and information about breathing is communicated to the cerebellum via CO2 receptors in the brain stem including the inferior olive/climbing fibre system and cerebellar nuclei (Band et al. 1980; Nattie 1999; Nattie et al. 1995; Oyamada et al. 1998; Pineda and Aghajanian 1997; Wise et al. 2004; Xu and Frazier 2000; Xu et al. 1994; Xu et al. 2001; Zhang et al. 1998). The next time you watch Usain Bolt charging down a 100-meter track to secure a gold medal be assured that on that trial the neocortex was not being used during task execution—even though without a neocortex Bolt could never have achieved such high-calibre performance (Hebb 1949; Vanderwolf 2007).

Footnote 1: Casual breathing (rather than exerted breathing) has been associated with neocortical rhythmicity (Zelano, Gottfried et al. 2016). This suggests that learning, a process mediated by the neocortex (Hebb 1949), is optimal under calm conditions rather than under violent conditions. So, the next time you think that beating knowledge into the head of your child is a competitive strategy, change your strategy.

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