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Fig. S1 (Savic & Lindström, 2008). Part of the left cerebral hemisphere VOI in a male heterosexual subject. Subject’s right side is to the right in the image.
Since PNAS is not, to my knowledge, a true peer-reviewed journal. I've decided to post a little review of the article addressing its validity.
[NOTE: that is not entirely true. Read the footnote1 for an explanation.]
Shusaku continues with this biting critique:
1) Structural MRI measruements - the regions of interest (ROIs) demarcated in this study included the ventricles, which are not gray or white matter. This indicates that the measurements of structural asymmetry can easily be confounded by differences in ventricular size. In which case, it is equally likely gray or white matter asymmetry cannot be determined by the results presented. There result of differences in structural asymmetry are further undermined by the fact that the cerebellum demarcations only contain gray/white matter and show no differences in asymmetry between subject groups.
Among studies explicitly comparing the entire hemispheres most, although not all, suggest that the right hemisphere is larger in men (32, 34, 38, 39).
By contrast, in a study on sex differences in cortical thickness, Sowell et al. (2007) opined:
Given the apparent specificity of differences in male and female cognitive advantages, and regional specificity of brain–behavior relationships, global differences in brain size between the sexes that have been readily observed with relatively gross methods might not be the most relevant structural dimorphism when investigating neural substrates of sex differences in cognition.
This is a very important point, because a major result in their study was that gray matter thickness in the right inferior parietal and posterior temporal cortices was 0.45 mm thicker in women than in men [all of unknown sexual orientation], as illustrated below.
Fig. 1 (Sowell et al., 2007). (A) Maps of differences between the sexes in thickness of gray matter (males coded 1, females coded 0 for all maps displayed) for the entire group of 176 subjects showing differences in gray matter (in millimeters) between the male and female subjects according to the color bar on the right. Warmer colors (less than 0 on the color bar) are regions where gray matter thickness is greater in the female than in the male subjects, and cooler colors (greater than 0) are regions where the males have greater gray matter thickness than the female subjects. Note the approximately 0.45 mm increase in cortical thickness in females in the right posterior temporal lobe.
And that's only by way of illustrating Shusaku 's point #1contra Savic and Lindström. He also criticized the amygdala functional connectivity analysis (a whole 'nother topic), the assumption that learning and environment cannot account for the results, and the exclusion of bisexuals (although all studies seem to exclude them, so that one's not really fair).
There are many more technical problems with the analytical techniques used in this study. I'm not going to bother going through all of them. Needless to say, this study proves absolutely nothing, and is just another example of bad science. As an MRI researcher, this article offends me personally, for it gives MRI and PET research a bad rap; I'm going to go throw up now.
Footnote
1 There are multiple routes to publishing in PNAS. One track is direct submission to the editors, who send it out for peer review. Another track is for a member of the National Academy of Sciences to either sponsor ("communicate") a paper by other authors or to submit ("contribute") his/her own paper (both without peer review from the PNAS Editorial Board). Savic and Lindström submitted to the Editorial Board, so the paper did go out for peer review. However, I noticed something that's not usually allowed, i.e., the acting Editor for the manuscript is affiliated with the same institution as the authors (Karolinska Institutet).
References
Savic I, Lindström P. (2008). PET and MRI show differences in cerebral asymmetry and functional connectivity between homo- and heterosexual subjects. Proc Natl Acad Sci. June 16, 2008, 10.1073/pnas.0801566105.
Sowell ER, Peterson BS, Kan E, Woods RP, Yoshii J, Bansal R, Xu D, Zhu H, Thompson PM, Toga AW. (2007). Sex Differences in Cortical Thickness Mapped in 176 Healthy Individuals between 7 and 87 Years of Age. Cerebral Cortex, 17(7), 1550-1560. DOI: 10.1093/cercor/bhl066.
Harold Mouras, at University of Picardie Jules Verne in Amiens, France, and his colleagues wanted to understand the cerebral underpinnings of visually-induced erections.They suspected there might be a role for mirror neurons, a special class of brain cell that fires both when people perform an action and when they observe it being performed.The researchers invited eight young men into the lab and asked them to view three types of video clips. Along with late-night fishing documentaries and snippets of Mr Bean, the volunteers got to see erotic videos of [XXX]...1
This isn't the first study to invoke the specter of mirror neurons as a critical aspect of responsiveness to viewing porn. Ponsetti et al. (2006) showed pictures of male and female sexually aroused genitals to gay and straight male and female participants [see An "Endophenotype" For Sexual Orientation? for a full description of that study]. The authors summarized their results as follows:
Consistent with our prediction, the ventral striatum and the centromedian thalamus, showed a stronger neuronal response to preferred relative to non-preferred stimuli. Likewise, the ventral premotor cortex which is a key structure for imitative (mirror neurons) and tool-related (canonical neurons) actions showed a bilateral sexual preference-specific activation, suggesting that viewing sexually aroused genitals of the preferred sex triggers action representations of sexual behavior.
At the time, I said:
Here's a thought. According to Wikipedia,
a mirror neuron is a neuron which fires both when an animal performs an action and when the animal observes the same action performed by another (especially conspecific) animal. Thus, the neuron "mirrors" the behavior of another animal, as though the observer were himself performing the action.
The Neurocritic is as skeptical as anyone about the mirror neuron craze, but if the PMv activity in this experiment is really imitative in nature, or even "empathetic" (instead of motor imagery or motor preparation), then wouldn't same-sex genitals elicit greater activity than opposite-sex genitals, regardless of sexual orientation?
To investigate the hypothesis that the activation of the mirror-neuron system could be part of the neural mechanisms regulating visually-induced sexual arousal, including the erectile response, we examined whether the response of the mirror-neuron system to sexually stimulating video clips is correlated with the erectile response of healthy volunteers.
The participants were ten healthy heterosexual males somewhere between the ages of 18 and 60. The authors hypothesized that
a neural pathway linking the mirror-neuron system to neural structures controlling erection could be represented by the efference of the frontal operculum – a region containing mirror neurons - to the insula.2
The ventral premotor cortex in Brodmann area 6 is posterior to the frontal operculum (Tomassini et al., 2007). Do we really know that the frontal operculum contains mirror neurons? The most [only]definitive evidence for mirror neuron-type activity is from single-unit recording, not from fMRI. Do monkeys even have a frontal operculum? Yes, but it seems to be mostly gustatory.
Vilayanur Ramachandran, at the University of California at San Diego, who also studies mirror neurons, calls it a "bold" study, and congratulates the group on defying the taboo on studying human sexual physiology.While he thinks it is perfectly plausible that mirror neurons play a role in how porn turns us on, he says more needs to be done to understand what that role is. For a start, he says, a large number of the brain's structures seem to be involved, not just the pars opercularis, and the interaction between these regions in response to porn is unclear. "It doesn't give you an experimental lever into the problem," he adds.And while Ramachandran agrees that the timing of mirror neuron activation and erection is probably critical, fMRI isn't accurate enough to show clearly what is going on with these brain regions over such short time frames.
Ramachandran isn't usually one to show restraint in interpreting data, but here he's right that the BOLD signal was correlated positively with the plethysmographic signal in multitude of brain regions. The authors focus on Frontal operculum, Precentral gyrus, Middle frontal gyrus, Postcentral gyrus, Inferior parietal lobule, Postcentral sulcus, Supramarginal gyrus, Anterior insula, and Posterior insula in the main body of the text, but one can...
(see exhaustive list of regions in Tables 1 and 2 of the online electronic supplementary material)
...once they're online (they're not yet).
ADDENDUM: My criticism of this statement, "Pars opercularis (Brodmann area [BA] 44) is a likely homologue of a subdivision of area F5 of monkeys" appears to be justified. In a recent review of 24 fMRI studies, Morin and Grèzes (2008) discovered that
Observing biological actions with a physical target, compared to a visual control showing no action at all, consistently activated the ventral premotor cortex (BA 6), and did so significantly more than observing target-less actions (with the same control). In contrast, the activity in BA 44 ("Broca’s area") was not modulated by the presence or absence of targets. We propose that the ventral precentral gyrus, and not BA 44, shares the visual properties of "mirror" neurons found in area F5 of the macaque brain.
Footnotes
1 Edited to avoid search engine hits to this blog from terms like "s*roking n**ed women, enjoying f***atio and engaging in interco**se."
MOURAS H, STOLERU S, MOULIER V, PELEGRINI-ISSAC M, ROUXEL R, GRANDJEAN B, GLUTRON D, BITTOUN J. (2008). Activation of mirror-neuron system by erotic video clips predicts degree of induced erection: an fMRI study. NeuroImage DOI: 10.1016/j.neuroimage.2008.05.051. Although visually-induced erection is a common occurrence in human male behaviour, the cerebral underpinnings of this response are not well-known. We hypothesized that the magnitude of induced erection would be linearly correlated with the activation of the mirror-neuron system in response to sexually explicit films. When presented with sexual video clips, eight out of ten healthy subjects had an erectile response demonstrated through volumetric penile plethysmography. The level of activation of the left frontal operculum and of the inferior parietal lobules, areas which contain mirror neurons, predicted the magnitude of the erectile response. These results suggest that the response of the mirror-neuron system may not only code for the motor correlates of observed actions, but also for autonomic correlates of these actions.
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While the results were striking, they would be more convincing if the authors had matched the groups for IQ, education and measures of depression and anxiety, said Suzanne Corkin, a professor of behavioral neuroscience at the Massachusetts Institute of Technology, in an e-mailed statement. Also, the authors are "overly dismissive'' of the potential role of environmental influences, Corkin said.
"In short, I would be reluctant to draw strong conclusions about heterosexual versus homosexual brain structure and connectivity from this single experiment,'' Corkin said. She wasn't involved in the study.
That quote came from an article written by Elizabeth Lopatto, which is commendable for interviewing experts and providing background on the subject. Read the whole article (excerpt below).
June 16 (Bloomberg) -- Gay men and straight women share brain characteristics that suggest sexual preferences may be innate rather than learned, researchers said. Lesbians and heterosexual men also had similar brain tendencies.
A study of 90 adults showed similarities between gay men and straight women in a part of the brain linked to emotional response called the amygdala, and a similar finding for lesbians and straight men. The research also found lesbians and heterosexual men had larger right brains, the side associated with spatial ability, while the left and right brains of both gay men and straight women were more symmetrical.
The study, published in the Proceedings of the National Academy of Sciences, adds to research that suggests a biological basis for homosexuality, researchers said. Earlier studies have mostly focused on behavioral differences and similarities.
Reference
Savic I, Lindström P. (2008). PET and MRI show differences in cerebral asymmetry and functional connectivity between homo- and heterosexual subjects. PNAS. June 16, 2008, 10.1073/pnas.0801566105.
‘...how narrow [banausisch] a matter it would be if there were cells of morality and immorality, such as virtue cells, murder cells, or cells responsible for rage. Things will surely be more complicated.’ -German psychiatrist Julius Koch (1894: 40–1), as quoted by Verplaetse (2004)
Those who cannot remember the past are condemned to repeat it.-George Santayana, The Life of Reason, Volume 1, 1905
Children of Uganda supports two orphanages in Uganda, as well as children living with HIV-positive widowed mothers, and has over 600 children under its care.
The localization of morality was discussed with equal reserve. Criminal anthropologists who looked eagerly for neurological and biological mechanisms to explain human immorality and criminal behaviour unanimously denied the existence of an isolated moral centre or moral organ that might be pathologically damaged. The French criminal anthropologists Gabriel Tarde and Alexandre Lacassagne dismissed the idea of a localizable moral sense as impossible or even ridiculous (Lacassagne, 1908: vii; Tarde, 1899: 240). ... Hans Kurella firmly asked that criminal anthropology should entirely abandon old-fashioned concepts such as an innate conscience, whether it was thought of as a localizable brain centre or not (Kurella, 1893: 204).
But really, the occipital lobes? Where did he get that idea? Verplaetse (2004) continues, quoting Benedikt:
When I freed the first brain (the brain of a robbing murderer) from the cranial cavity, his crime became clear to me with an unprecedentedly anatomical transparency. The occipital lobes did not cover the cerebellum and in this discovery I discerned the crucial distinction between man and animal.
OK, then. Onward and upward, to 2008 and the neural encoding of distributive justice.
...researchers at the California Institute of Technology have discovered that reason struggles with emotion to find equitable solutions, and have pinpointed the region of the brain where this takes place. The concept of fairness, they found, is processed in the insular cortex, or insula, which is also the seat of emotional reactions.
"The fact that the brain has such a robust response to unfairness suggests that sensing unfairness is a basic evolved capacity," notes Steven Quartz, an associate professor of philosophy at Caltech and author of the study, voicing a sentiment that anyone who has seen children fight over a treat can relate to.
"The movement to look into the neural basis for ethical decision making is only about seven years old," Quartz adds. "This is the first study where people made real decisions with real consequences."
So the insula is the seat of emotional reactions!! And investigations of the neural basis for ethical decision making are only 7 years old!
According to Wikipedia, the insula "lies deep to the brain's lateral surface, within the lateral sulcus which separates the temporal lobe and inferior parietal cortex. These overlying cortical areas are known as opercula (meaning "lids"), and parts of the frontal, temporal and parietal lobes form opercula over the insula."
It's a pretty large area. Besides being crowned the "seat of emotional reactions" (whatever that means), portions of the insula have been associated with interoceptive awareness, visceral sensation, pain, autonomic control, and taste, among other things... a lot of other things. Do a search of the BrainMap database using just two of the many insular foci reported by the Caltech researchers and you'll see activations related to action execution, speech, attention, language, explicit memory, working memory, and audition.
...how goods and benefits should be dispersed throughout a society in a fair and just manner. As an extreme example of this dilemma, imagine you are commissioned to deliver 100 lbs. of food to a famine-stricken region that consists of two villages a hundred miles apart. If you deliver half of the food to the first village, then travel to the second, 30 lbs. of the food will spoil during the trip...
Philosophers have offered several solutions to debates of this nature. Utilitarianism... asserts that one’s primary goal should be the achievement of a maximal amount of good or happiness. In the situation described above, a utilitarian might opt to deliver all of the food to the first village. ..... Another approach to such a quandary is known as deontological ethics, which emphasizes not the consequences of one’s actions, but whether the actions are right or wrong, just or unjust. From a deontological perspective, it would be unjust to distribute the food unequally.
To examine the trade-off between equity and efficiency, Hsu et al. (2008) devised a task in which the participants decided on how to allocate money to children living in an orphanage in northern Uganda.
In each trial, participants decided whether varying allocations of money, denominated in meals, would be taken away from either of two groups of children; the participant’s choice was to decide from whom to take.
You can see an example of an experimental trial in this must-see movie from the paper's Supporting Online Material (embedded below).
Movie s2 No Switch Trial. Illustration of a trial where the subject does not switch the lever. Animation speed is increased for illustration purposes. See Fig. 1 for actual duration of events and screens.
During the Switch, insular activity was correlated with the level of inequity, but so was activity in the right postcentral gyrus and the medial frontal gyrus (Table S12). Before the Switch (during the Display), inequity correlated with insular activity and with activity in R Brodmann area 10, L inferior temporal gyrus, R medial frontal gyrus, L posterior cingulate, L BA 39, R superior temporal gyrus, R precuneus (Table S11)... I needn't go on.
The authors conclude:
Against utilitarianism, our results support the deontological intuition that a sense of fairness is fundamental to distributive justice but, as suggested by moral sentimentalists, is rooted in emotional processing. More generally, emotional responses related to norm violations may underlie individual differences in equity considerations and adherence to ethical rules.
...rationalization of inequality statistically mediates the relationship between conservatism and happiness. In other words, it suggests that at least part of the reason conservatives are happier than liberals is that they're more likely to rationalize inequality.
But did we need fMRI to tell us that? Is it really all in the insula? Couldn't we have more or less learned the same thing by obtaining peripheral autonomic measures, like heart rate, blood pressure, and skin conductance?
Footnote
1At least, if not earlier. Any historians of neuroscience out there?
References
Hsu M, Anen C, Quartz SR. (2008). The Right and the Good: Distributive Justice and Neural Encoding of Equity and Efficiency. Science, 320(5879), 1092-1095. DOI: 10.1126/science.1153651
Koch J. (1894) Die Frage nach dem geborenen Verbrecher (Ravensburg: Otto Maier).
Verplaetse J. (2004). Moritz Benedikt's (1835-1920) Localization of Morality in the Occipital Lobes: Origin and Background of a Controversial Hypothesis. History of Psychiatry, 15(3), 305-328. DOI: 10.1177/0957154X04039354
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