Brain evolution and PFC fractionation Flashcards
Brain volume
Steadily increased relative to body weight in homo-lineage by a factor of approx. 2 from 2 million years ago.
(Lewin & Foley, 2004).
Human brain no bigger than it should be
Reached current size approx. 100,000 years ago.
Symbolic developments occurred 50k to 40k years ago.
Brain size is not everything.
- among humans, only small proportion of intelligence differences attributed to size. (Rushton & Ankey, 2009).
Paleoneuroanatomical evidence
Hominin cranial fossils preserve evidence of:
- overall brain size.
- cerebral asymmetry.
- cortical sulcul patterns - leaves impressions on the endocra surface.
Fossils suggest 3 major stages of hominin brain evolution
Stage 1 (3.5-2mya):
- brain reorganisation without substantial expansion - includes relative expansion of posterior parietal association cortex at the expense of the occipital cortex.
- may have been important for emergence of stone tool making by 2.6mya (Semaw et al, 2003).
- some suggest stage 1 involved in prefrontal lobe shape (Falk et al 2000).
Stage 2 (2-0.5mya):
- sudden increase in brain size associated with appearance of homohabilis.
- followed by gradual expansion related to body size increases in homo-erectus.
- first appearance of modern human like cerebral asymmetries in hobo habilis including enlargement of the Broca’s cap region in left LPFC (BA44).
Stage 3 (0.5-0.02mya): - past 15k years decreasing body size brought human mean brain size down a bit.
Overall (excluding Broca’s area cap enlarging) evidence of frontal lobe size and reorganisation is limited.
- fractionated function.
Evidence from comparative neuroanatomy: relative size of the frontal cortex?
Differences are large in size of FC between humans and primates.
- but FC in humans and great apes occupies a similar proportion of the cortex of the cerebral hemispheres (Semendeferi, Schenker & Damasio, 2002).
Evidence from comparative neuroanatomy: frontal vs. prefrontal
Not relative size of the frontal cortex but of the PFC - evidence of PFC reorganisation.
Prefrontal area argued to be fractionated (a substance is divided during phase transition into smaller quantities). in 1 of 3 ways:
- presence of granular layer 4 (Stellate and other smaller cells).
- projection area of the mediodorsal nucleus of the thalamus.
- motorically “silent” area when stimulated.
Evidence from comparative neuroanatomy: larger in humans
Human PFC especially enlarged compared to great apes - Passingham & Smaers (2014).
BA10 (Polar PFC) is larger in humans relative to the rest of the brain than in apes - Semendeferi et al (2001).
Cytoarchitectonics & granular cells of the PFC: granule cells
= interneuron.
- defined by its smallness.
Benefits of size:
- density and number of connections.
In the PFC: only primates have granular layer 4 in PFC.
- thickness of layer 4 increases as one foes from caudal to rostral along the medial and orbital surfaces of the frontal lobe.
- area can be agranular enough to warrant exclusion from special status even if some granule cells present.
Cytoarchitectonics & granular cells of the PFC: cerebral cortex layers
Cerebral cortex = outer layer of cerebrum/cortex.
- largest and most prominent part of the brain.
- cerebral cortex has 4 lobes.
Layers:
- molecular layer.
- external granular layer.
- external pyramidal layer.
- internal granular layer.
- internal pyramidal layer.
- multiform layer.
Cytoarchitectonics & granular cells of the PFC: phases of evolution of the PFC
Early mammals develop agranular areas of PFC: medial and orbital PFC/insular cortex.
- primates alone have granular cortex.
- rats only have agranular PFC.
Lateral and polar granular PFC last to appear during anthropoid evolution.
Granular PFC appeared in early primates as they adapted to life confined by trees - caudal PFC and areas of OPFC:
- function in the assessment of value or primary reward (e.g. food - Passingham & Wise, 2012).
Several new granular PFC areas appeared during anthropoid evolution - grew larger, foraged more (by reducing choices that increased risk of predation or wasted effort), became dependent on food and vulnerable to falls:
- dorsal PFC (BA9/46).
- ventral PFC (BA45/47).
- polar PFC (BA10).
Lateral and polar granular PFC - last to appear during anthropoid evolution.
Brain became wider and more rounded at the front during homonoid evolution.
Cytoarchitectonics & granular cells of the PFC: what did granular PFC add?
Evolved to implement new, faster, general-purpose mechanism - in response to adaptive pressures.
- supports older, reinforcement-learning mechanism.
Granular PFC generates goals appropriate to goals and needs.
Cytoarchitectonics & granular cells of the PFC: dorsal paracingulate cortex connects to dorsal PFC
Modern humans probably evolved after Broca’s expansion seen in homo habilis.
- when frontal lobe PFC reached modern state - before advanced tool use, abstract thought, language etc (approx. 70-40kya).
Elston et al (2006): granular cortex is 80% in humans and 55% in chimps.
Elston et al (2001): pyramidal cells in layer 3 = 70% more spinous in humans then monkeys.
Schenker et al (2005): human PFC has larger volume of short nerve fibre connections connecting parts of PFC.
Areas of human specialisation (at cell level of PFC):
- broca’s area (BA 44/45).
- lateral part of the polar PFC (BA 10).
- dorsal anterior cingulate/medial PFC (BA 32).
> BA 10 and BA 32 lack homologues in monkeys.
Cytoarchitectonics & granular cells of the PFC: connections
PFC = brains controller.
- has connections.
Synaptogenesis last longer in PFC =than other regions (Bianchi et al, 2013; Levitt, 2003).
Cytoarchitectonics & granular cells of the PFC: brain development, mirrors brain evolution
Cortical expansion during evolution matches expansion during development (Hill et al, 2010).
Ultimately, newer PFC regions (lateral and polar) are better learning devices (Passingham & Wise, 2012).
Leaves us more open to cultural influences than other species whose brain stops developing earlier and are more influenced by genes.
Cytoarchitectonics & granular cells of the PFC: abstraction increases with granularity
Processing hierarchy in brain (Badre, 2008):
- posterior
> anterior
> caudal
> rostral.
- processing moves from concrete to abstract.
Granular PFC at the apex of processing hierarchy (Passingham & Wise):
- allowing integration of all info necessary to generate goals from current context and events based on knowledge of current value.
Cytoarchitectonics & granular cells of the PFC: PFC summary
Brain got wider and more round at front.
- lateral and polar regions (populated by granular cells) - during hominoid evolution.
Regions of PFC enlarged compared to other apes:
- particularly BA10.
Evidence of frontal lobe changes during hominin evolution:
- particularly in Broca’s cap in lateral PFC which would have widened brain further.