The Neuroanatomy of Homosexuality

Spring 2010 Vol 1 Issue 2 thenerve

The Religious Brain

Neuroscience and the Military

The Rise of the Cyborgs Mind and Brain Society

The Mind and Brain Society (MBS; formerly known as the BU Organization for the Mind and Brain Sciences) was founded in the fall of 2008 in concert with BU’s new Undergraduate Program in Neuroscience. The group aims to create a network for undergraduate students who wish to take an active role in current issues and research. MBS serves as a hub for not only Neuroscience majors, but all students interested in Psychology, Biology, Philosophy, Computer Science, etc. Our goal is to support an ea- ger multidisciplinary undergraduate community with the conversations and resources fundamental to Neuroscience today. Throughout the academic year, MBS hosts events spotlighting many different fac- ets of Neuroscience. We hold discussion sessions during which we informally discuss a topic of interest over coffee; previous topics include “The Neuroscience of Religion” and “NeuroEthics.” The group also hosts research presentations by BU professors and screenings of thought-provoking films containing neuroscience motifs. Cover: Brainbow mouse hippocampus. Image by Tamily Weissman. The Brainbow mouse was produced by Livet J, Weissman TA, Kang H, Draft RW, Lu J, Bennis RA, Sanes JR, Lichtman JW. Nature (2007) 450:56-62. CONTENTS Spring 2010 Vol. 1 Issue 2

RESEARCH IN BRIEF 6

ARTICLES The Rise of the Cyborgs by Darrien Garay 12

Brain Battles: Neuroscience and the Military by Monika Chitre 15

The Religious Brain by Pinkey Shah 19

The Special K Challenge for Depression by Aisha Sohail and Jeff Wessell23

REVIEWS Recognition Memory: One process or two? by Danielle Miller 27

The Neuroanatomy of Homosexuality by Claire Bryson 33

Huemer’s Theory of Perception: Analysis and Objections by Ethan Rubin 37

OPINION Homemade Terror by Darrien Garay 46

SPRING 2010 | 3 FROM THE EDITORS

he U.S. media has been consumed by healthcare Editors-In-Chief reform for the past year or so, arriving at a climax Grigori Guitchounts Twith the historic vote in the House of Representa- Kimberly LeVine tives on March 21. Whether or not the new legislation will Editors alleviate the country’s problems remains to be seen. No Neil Datta matter what your opinion on the changes making their way Doug Hidlay through America’s healthcare system, we cannot deny that they will have an impact on our community. Our hopes are Associate Editors that a new system will only enhance the neurological care Lauren Joseph for those around us. Here at The Nerve, we hope you will Monika Chitre take a few minutes to forget about the politics and enjoy the Natalie Banacos science. Jennifer Richardson Having set out to conquer the world by means of study- Frank DeVita ing abroad, our editorial team brings you the Second Issue Artwork from three continents, including Europe and South America. Kayla Ritchie Our goal was to collect articles as far-reaching as our staff. From the relationship of the military with neuroscience to MBS Staff Darrien Garay the neuroanatomy of homosexuality, we are confident you Megan Mataga willThis find andsomething other issues inside of to The stimulate Nerve wouldyour brain. not have been Macayla Donegan possible without the generous help of Howard Eichenbaum, Shea Gillet Paul Lipton, Lindsey Clarkson, Denise Parisi, Zachary Bos, Meghna Majithia and Jarret Frank. Advisors Paul Lipton - Grigori Guitchounts and Kimberly LeVine Zachary Bos Editors-In-Chief

4 | THE NERVE

RESEARCH IN BRIEF Vast Parallel Processing in a Retinal Interneuron to support cellular depolarization in vivo. As such, this study’s ini- tial morphological data suggesting independent processing functions is lending

studies into calcium’s activity significant credence. Further even though transient in- tensewithin Ca2+ A17 activity cells shows can result that, in GABA release, even studies stimulating multiple Ca2+

neurite could not experimen- tallychannel propagate patches any along electronic an A17 information beyond other The eye is a truly wondrous processing organ synaptic contacts in the immediate vicinity, and only composed of six primary cell types separated into two weakly even then. primary plexiform layers. While rods and cones, the actual light responsive cell types, are well character- 150 Given primarily that independentthese A17 cells synapses, are able theto receive activity and of more complex in their activity and ability to modulate thisprovide type RBC of amacrine contact-specific cell mimics feedback the atdendritic greater intethan- visualized, horizontal input. In and their amacrine study, Grimes cells are et al. significantly examined gration seen in cortical processing interneurons. The morphological, biophysical, and synaptic properties for granted in upper cognitive pathways seems to be asone related specific to type their of amacrine feedback cells, onto the rod A17, bipolar and their cells mirroredfine-tuning even that within has been the essentials studied and of the may visual be takenpath- (RBCs); ultimately, concluding that, at least for the way. This phenomena likely accounts for some of the hundreds of parallel microcircuit feedback loops in stimulate the RBCs and the ability to perceive subtle A17, amacrine cells are uniquely structured to create- changesfine visual in acuity light stimulusseen under even low-light when conditions the magnitude that ing on the output of bipolar cells and the overall rod of stimulus is considerably lower than normal visual pathway,order to provide which isan responsible incredibly forfine visionlevel of under process low parameters. Additionally, by condensing all the mul- ambient light. titude of microfeedback circuits into a single cell, the - visual system reduces the “wiring cost” necessary to ing in combination with cell modeling studies sug- Both fluorescent and electron microscope imag light rod pathway. The Grimes et al. study has laid im- cell extending into the layer of RBCs is able to receive maintain the fine level of feedback needed in the low- synapticgested that output each long, and unbranching return appropriate neurite processingof the A17 responsibilities of the modulatory pathways that al- largely independent of the activity at any but the most lowportant for thegroundwork incredible for diversity finally piecingof signals out and the visual exact discrimination seen in the stimulus-transducing por- cell to reciprocally modify hundreds of inputs with tion of the optical pathway. limitedproximal overlap parts toof neighboringthe cell, allowing points for of synaptica single conA17- tact. Beyond these morphological features limiting — Doug Hidlay the ability of electronic conductance, examination of both sodium and potassium channel response to in- : Grimes WN et al. Retinal Parallel Processors: More than 100 Independent Microcir- do not cause more than a transient depolarization of cuitsoriginal Operate paper within a Single Interneuron. Neuron. 65, localjected portions current ofin neuritesA17 cells in shows vitro, thatwith these little evidencechannels

873–885, 2010. 6 | THE NERVE RESEARCH IN BRIEF Mindblind Eyes in Asperger Syndrome It is well known that autism spectrum disorders ing movement and did not assign a false-belief to the frequently plague sufferers with impairments in com- character in the task. munication and social interaction. A popular hypoth- The results show that adults with Asperger syn- esis breaking into what underlies these impairments drome do not spontaneously predict others’ actions is that autistic children lack Theory of Mind (ToM), in a nonverbal situation. This impairment charac- the ability to assign mental states to oneself or others; terizes mindblindess in autism spectrum disorder that is to say, one’s ability to understand others’ feel- patients. Senju et al suggest that nonverbal mental ings or viewpoints based on the unique set of infor- state assignment skills require spontaneous encoding mation available to one. In the Sally-Anne false-belief of socially relevant information, a skill that individu- task (FBT), a popular test of the Theory of Mind (ToM) als with Asperger syndrome lack, which lends to dif- skill, children are asked to predict a girl’s actions when she looks for a marble that has been moved from its study proposes that it is possible to enhance explicit, usual location without her knowledge. Do children verbalficulty with mental social state interactions. assignment On through the other compensa hand, the- with Asperger syndrome understand Sally-Anne’s tory learning. mental state or are they unable to distinguish their own omniscience from Sally’s misinformed view? — Kimberly LeVine Those with Asperger are, in fact, able to correctly predict Sally’s action, confounding the mindblindess : Senju A, Southgate V, White S, and hypothesis supporting impairment in Theory of Mind. Frith U. Mindblind Eyes: An Absence of Spontaneous Senju et al propose that Asperger patients have Theoryoriginal of paperMind in Asperger Syndrome. Science the ability to reason through false belief scenarios but nevertheless maintain an impairment as- . 325, signing mental states. The study contrasts 883-885. 2009. results between verbal and nonverbal forms of the false-belief task to trace the divergence of mindblindess between typically developing participants and those with Asperger Syndrome. An earlier study by Southgate et al, using a paradigm similar to the Sally-Ann FBT, sheds light on the tendency of infants to not only look longer at unexpected events than at expected ones, but to make eye-movements toward expected events before they occur. Senju et al used this information to assess whether adults with Asperger syndrome can successfully assign a false belief as evi- denced by anticipatory looking. As expected, neurotypical (i.e. those without Asperger’s) adults and those with Asperger syndrome performed equally on the verbally instructed ToM tests (such as the Sally-Anne task). However on nonverbal ToM tests which measured looking bias, the Asperger group did toward the correct target as did neurotypi- calnot adults. show theThe sameAsperger significant group’s levellooking of bias did not differ from zero, supporting the theory that they did not anticipate the impend-

SPRING 2010 | 7 RESEARCH IN BRIEF

- Methylated Memories - sinegenerally residues have in difficulty fear-conditioned creating new mice, ones. the By study look at Memory is certainly one of the greatest myster- theing atUniversity the methylation of Alabama of H3 explored histones the on effects various of exly- ies of neuroscience. How do we store information perience on epigenetics in the hippocampus during for long periods of time? How do we learn? Recent memory formation. results published by Swati Gupta et al. of the Depart- The mice used in the experiments were all young ment of Neurobiology at the University of Alabama at adult males of the same species. One experimen- Birmingham provide a good start to answering these tal group underwent a process called fear-learning. questions. The study illuminates the role of genetics These mice were brought into the laboratory thirty in neuroscience: regulation of gene expression proves minutes before being placed in a training chamber. Af- important in the formation of long-term memories. ter two minutes in the chamber, they received electric Epigenetics is the regulation of gene expression shocks to their feet paired with a sound. The shocks based on factors other than alterations in a DNA se- occurred in sets of three, every two minutes. A second - experimental group was content-exposed, meaning ronmental factors. Effects often occur after transla- that they were put in the chamber for seven minutes tion,quence: once this the regulation gene has been is largely transcribed reflective into of mRNA envi and did not receive any shocks. Upon re-exposure to to be “read” by ribosomes, and a polypeptide chain the chamber this group did not exhibit freezing be- has been assembled accordingly. Phosphate and ace- havior, indicating they had not developed a fear re- tyl groups are able to unwind DNA to expose histones, sponse to the chamber. A third group of mice were the proteins that DNA wraps around when it con- latent-inhibition plus fear-conditioned. Latent inhibi- denses to form chromatin. Once these histones are tion involves exposing a subject to a new situation for exposed, methylation can occur. Depending on which a long period of time (in this case, two hours) before amino acid residues the methyl groups attach to, tran- beginning fear-conditioning by introducing the un- scription can be either silenced or activated. Inducing conditioned stimulus, the foot shocks (as experienced or inhibiting methylation is how external factors can by the fear-conditioned animals). This group ensured regulate the expression of genes. that associative learning was taking place: when they Gupta’s study examined methylation in the hip- were reintroduced to the training chamber 24 hours pocampus of mice. This region of the brain is notable later, freezing behavior was observed even though for its contribution to the formation of long-term they were not shocked upon re-entering the chamber. memories: human patients with damage to this area A control group received no experimental treatment. After the mice were trained, they were watched for freezing behavior when presented with the conditioned stimulus: the training chamber and the sound paired with the shocks. Once behavioral testing was complete, the CA1 area was re- moved from the hippocampus of each mouse. Antibodies were used to separate DNA from chromatin, and PCR was used to amplify and analyze the samples.

8 | THE NERVE RESEARCH IN BRIEF

Samples that were observed an hour after condi- tone deacetylases), with sodium butyrate (NaB) was tioning took place showed an increase in transcrip- other words, if the histones were lacking in acetyl had been contextually fear-conditioned as opposed groupsfound to because decrease of methylation HDAC suppression, on H3K9 less histones. methyla In- totion-activating the mice who H3K4 had trimethylationonly been context-conditioned. in the mice that tion occurred: a link between hippocampal acetyla- This observation links the methylation to associative tion and methylation. shown increased in both the contextual fear-condi- certain promoter regions on particular genes, exhib- tionedlearning mice processes. as well The as H3K9 in the dimethylation context-conditioned that was itingH3K4 the regulationmethylation of increased methylation specifically involved around in fear mice an hour after training. This transcription-silenc- conditioning. In general, fear conditioning tended ing methylation must be important to fear learning to demonstrate an increase in methylated DNA. This as well as context learning. In looking at the mouse methylation seems to serve as a signal for gene tran- brains 24 hours after training, it was found that both scription. kinds of methylation are reversible: amounts of meth- So how is all this genetic information useful to ylation decreased. our brains? According to Gupta’s paper, studies have shown that histone methylation levels in the hippo- that methylation of histones aids long-term memory campus may be able to help doctors identify schizo- formationAnother is findingthe importance that is supportive of two genes, of the Mll idea and phrenia earlier. Demethylase enzymes are being iso- eed. These two genes are essential to the synthesis lated, and their inhibition or upregulation could be used as treatments for many neurologic diseases. Epi- methylates this particular histone. Heterozygote Mll genetics is proving to be a useful tool in deciphering miceof H3K4-specific showed stronger methyltransferase, freezing behavior an enzyme than that eed - don’t! transferase and eed is to methyltransferases in gener- how our brains work, and how to fix them when they al.mice: Mll thisis particularly is because essential Mll is specific to the tomethylation H3K4 methyl pro- — Natalie Banacos cess observed. As long as mice were heterozygous or homozygous dominant for Mll and eed, they learned : Gupta S et al. Histone Methylation and remembered normally. Regulates Memory Formation. The Journal of Neuro- In another experiment, inhibition of HDAC (his- scienceoriginal paper

. 30(10):3589 –3599. 2010. homeostasis; thus, the polymor- Decreased Risk of Dementia is phism results in a lower concentra- tion in serum, increased HDL lev- Associated with CETP Polymorphism els, and HDL/LDL particle sizes. All of these are posited to play a role in Dementia is debilitating to phism in cholesteryl ester transfer cardiovascular protection. those affected; it is a major char- protein (CETP), which may be as- In the Ashkenazi population, acteristic of many neurological dis- sociated with a lesser and slower researchers have also found the eases, such as Alzheimer’s disease. memory loss/decline as well as a homozygous polymorphism at a In fact, the prevalence of demen- lower risk of dementia. tia is increasing as the population Previously, other researchers old individuals without dementia ages, resulting in public health and have documented that the CETP compared5-fold higher to ratethose in with75- to dementia. 85-year- economic burdens. However, de- polymorphisms has been associat- These studies purport a protective mentia does not affect all patients ed with longevity and lower cardio- association between these CETP in the same manner. In fact, some vascular risk, especially in samples polymorphisms and decreased de- patients enjoy a slower or lesser of Ashkenazi Jews that had higher mentia. However, CETP polymor- cognitive decline than others, much percentages of centenarians with phism studies on more general of it determined by genetics. Sand- the polymorphism than controls. populations have not previously ers, Wang , Katz, and colleagues CETP is a protein that is involved in demonstrated any cognitive bene- have investigated a gene polymor- central nervous system cholesterol

fits, possibly because of the limited SPRING 2010 | 9 RESEARCH IN BRIEF time frame they encompassed. risk for dementia or Alzheimer’s gotes also had lower risk of both Thus, Sanders, et al. looked Disease. dementia and AD. at the single-nucleotide polymor- Using the Free and Cued Se- These results may help sci- phism (SNP) at CETP codon 405, lective Reminding Test (FCSRT), where an isoleucine is changed to Sanders and colleagues assessed possible venues for treatment (i.e. a valine, and focused on the cogni- memory loss over time. The FCSRT CETPentists inhibitors, and doctors etc), not but only also find it tive effects over time. To test their is a test where learning conditions may lead to greater understand- hypothesis that this polymorphism are controlled by having subjects ing of the etiology of dementia. Al- would be associated with lower search a card containing four pic- zheimer’s may soon be more treat- memory decline and lower risk of tures of items that correspond to able than previously anticipated. dementia, they conducted a longi- unique category cues. After these tudinal prospective cohort study — Neil Datta test immediate cued recall and community-dwelling adults with- eventuallyitems are conduct identified, three researchers test trials : Sanders AE, et al. outthat dementia incorporated at baseline, 70+-year-old who consisting of free recall, followed Association of a functional poly- were registered with the Einstein by cued recall. Sanders and col- morphismoriginal paper in the cholesteryl ester Aging Study. In it, they investigated leagues ascertained that memory transfer protein (CETP) gene with associations between the CETP declined in polymorphism homo- memory decline and incidence of polymorphism genotype and longi- zygotes by a rate 51% slower than dementia. JAMA tudinal memory performance and in the control group. The homozy- 2010. . 303(2):150-158. Neuronal Transplantation Induces does nothing to hinder vision or reorganize cortex; animals wearing eye patches for several days can still Plasticity in Visual Cortex see well after the patch is taken off. Southwell capital- Normal neural development features critical pe- ized on this fact to test whether neuronal transplan- riods during which maximal learning occurs in a sen- tation affects visual cortex during MD post critical sory modality or neural system. The most commonly studied critical periods are those for visual perception induced a shift in ocular dominance columns just as and language acquisition. The former was reported in itperiod. does duringIndeed, the MD regular 33-35 criticaldays after period. transplantation Transplan- tation had the strongest effect when the donor cells performed a series of groundbreaking experiments reached the age at which the regular critical period thatthe 1960’s earned by them David the HubelNobel Prizeand Torsten in medicine. Wiesel, who occurs, regardless of the host’s age. During the critical period of visual perception, Furthermore, the transplanted cells migrated both eyes must be open and receiving meaningful in- into all layers of host visual cortex and developed put in order to be functional later in life. Occlusion of morphologies of mature inhibitory neurons with re- one eye (Monocular Deprivation, MD) during the criti- ciprocal synaptic connections. And even though trans- cal period induces rapid reorganization in visual cor- planted neurons’ synaptic inputs were only one-third tex that leads to the takeover of cortical ‘real estate’ as strong as those in native synapses, as measured by by the non-deprived eye’s axons, leaving the animal excitatory post-synaptic potentials, they were more blind in the occluded eye. Dogma has it that unsuc- numerous. cessful learning during the critical period (as in mon- - ocular deprivation) cannot be reversed later in life; erative medicine. Perhaps in the future, we will be once the animal is blind in the deprived eye, nothing able These to reverse results seemingly are promising permanent to the field neurological of regen conditions by introducing new cells into neural net- Derek Southwell and colleagues at UCSF show works and jump-starting neuroplasticity. thatmay fixthis that. is not necessarily the case. In a paper published in Science in February, they report that trans- — G. Guitchounts plantation of inhibitory neurons into mouse visual cortex after the critical period is over reactivates it, : Southwell DG, et al. Cortical Plastic- allowing vast reorganization. ity Induced by Inhibitory Neuron Transplantation. After the critical period, monocular deprivation Scienceoriginal paper

. 327: 1145-1148. 2010. 10 | THE NERVE RESEARCH IN BRIEF

Research finds neurons drive behavior

Daniel Huber and colleagues reported in a paper in Nature their research on the link between the ac- somatosensory cortex and perceptual and cognitive functionstivities of inspecific mice. Thegroups primary of neurons goal in conductingin the primary this research was to develop an alternative to the method of electrical microstimulation in order to isolate groups of neurons and study their activity in vivo. The drawback of electrical microstimulation is that the accuracy at which axons are excited is typically poor. Alternatively, through the use of photostimulation, in which a light-gated algal channel channel- rhodopsin-2 (ChR2) was introduced into the barrel cortex pyramidal neurons of mice, the discrimination between cells is far greater, and a considerably lower neurons was then determined, and it was found that number of neurons required stimulation in order to with decreasing light intensity, neurons abruptly produce robust behavior. ChR2 photostimulation in - vivo reliably generated stimulus-locked action poten- - tials at frequencies up to 50 Hz. An untrained group switched from firing action potentials with high prob of mice was capable of detecting brief trains of action preciselyability to timedfiring nospike action trains potentials in vivo. at all. It was con - firmedTo test that whether ChR2 can awake transduce mice could photostimuli learn to report into potentials (five light pulses, 1ms, 20 Hz). After train- in the barrel cortex, light pulses were delivered to rons.ing, however, Even fewer mice couldneurons detect (approximately a photostimulus 60) firingwere ChR2-GFP-expressingphotostimulation of layer neurons 2/3 in pyramidal freely moving neurons ani- a single action potential in approximately 300 neu mals. The mice were temporarily connected to an LED ms). controller and trained to associate photostimulation requiredTo conduct for longer this stimuli research, (five action in utero potentials, electropor 250- of CHR2-GFP expressing neurons with water reward tation was used to introduce ChR2 fused to a green on one of two choice ports. After four to seven training sessions, mice were capable of reliably report- - ing photostimulation by choosing the correct port calfluorescent pyramidal protein neurons. (GFP) The (ChR2-GFP) responses together of ChR2-GFP- with a in the presence or absence of stimulation, unlike the expressingred fluorescent neurons cytosolic to photostimulation marker (RFP) into in anaesthe neocorti- control mice without electroporated neurons, which tized mice were characterized, and recordings were performed at chance levels even after 25 training sessions. photon targeted loose-patch recordings. Photostimuli The results of these experiments demonstrate consistedgathered from of light red pulses fluorescent produced neurons by a using blue light- two- head (the researchers made sure the mice could not theythat photostimulationprovide a far more of accurate layer 2/3 method neurons of canstudying drive seemitting when diode the LED (LED, was 470 on nm)or off). glued At tomaximum the mouse’s light therobust circuit behavior. basis ofThese perception findings and are cognition significant in invivo. that intensities, about half of the patched red neurons responded reliably with at most one action potential. An — Lauren Joseph increase of the photostimulation duration beyond 10 ms did not reveal additional responsive neurons. : Daniel Huber et al. Sparse optical mi- The relation between photostimulus intensity crostimulation in barrel cortex drives learned behav- and the probability of spiking ChR2-GFP-expressing iororiginal in freely paper moving mice. Nature. 451:61-64. 2008.

SPRING 2010 | 11 ARTICLES

The Rise of the Cyborgs

By Darrien Garay cyborg Cyborg. It’s a word that calls to mind unteered for an experimental speech-research images of a RoboCop-like organism clank- program, a decision that would lead to the im- ing around in some far off future. However, plantation of mechanical devices directly into our notions about the reality of cyborgs may his brain1. be obsolete. Successful clinical research has taken giant leaps forward in the intimate fusing of brain and machine. Take for exam- When Brains and Machines Can Chat ple the story of Erik Ramsey. he device implanted within Erik’s brain is part of a brain-machine interface left himOn the riddled evening with of severe November internal 5, 1999 trau- T(BMI). BMI, also known as brain-com- ma.Erik Erik’swas involved injuries inresulted a traffic in accidenta stroke that puter interface or neural interface system, damaged his brainstem1. Due to this stroke, is an emerging branch of neuroscience that Erik developed a neurological condition seeks to overcome the “limitations of our bod- known as Locked-in Syndrome. Locked-in ies by forming a direct interaction between patients are complete- the brain and the outside ly paralyzed and can world.” move only their eyes2. “is it really BMIs3 can be divided However, they maintain into two major catego- conscious awareness, ries: those that induce can reason, and sense possible to desired brain activity or every pain or discom- those that record brain fort. activity. Erik’s story could liberate the Cochlear implants have ended when he are one example of a BMI was 16, essentially in- which induces desired carcerated within his mind from patterns of neural activ- own body, but techno- ity. A processor above the logical advances and outer ear receives incom- tremendous determi- the confines ing sounds and translates nation are writing a them into digital signals. new chapter, one of These signals are then hope. Five years after of the body?” transmitted through the the accident, Erik vol- skin and skull to a micro-

12 | THE NERVE ARTICLES “locked-in patients are completely paralyzed and can move only their eyes” electrode array implanted directly onto the Her brain now wirelessly connected to auditory nerve. The array stimulates the audi- the robot’s limbs, Idoya was again placed onto tory nerve, bypassing inner ear dysfunctions the treadmill. In front of the treadmill was that cause the deafness. According to the FDA, a screen that presented live video of CB for - Idoya to watch as she moved. ceived cochlear implant surgery worldwide4. As the treadmill was switched on, Idoya as ofBMIs April, of 2009; the second 188,000 class, people those have that record neural activity, are being tested as aug- CB came to life. His robotic legs moved in al- mentation systems for patients suffering from mostbegan perfectto walk synchrony- her motor toneurons her gait. fired As – and the various degrees of paralysis. These systems researchers had hoped, the neural signals re- circumvent limb paralysis, connecting the corded from Idoya’s brain orchestrated CB’s - strides. tors. The machine then executes the mental Idoya was even able to keep the robot commandsbrain’s output the fibers patient’s directly muscles to machine can no effec lon- walking when she had stopped walking her- ger follow. self. The visual feedback, provided by the But is it really possible to liberate the screen, allowed just the thought of walking to successfully control the robot’s limbs. “We have shown that you can take signals mind from the confines of the body? across the planet in the same time scale that a Monkey Think, Robot Do biological system works. Here the target hap- pens to be a robot. It could be a crane. Or any Nicolelis and his colleagues at tool of any size or magnitude. The body does Duke University were able to not have a monopoly for enacting the desires Dr.successfully harness a monkey’s of the brain,” said Dr. Nicolelis6. brain impulses to command a robot, half a world away, to walk5. First, researchers implanted recording electrodes into the motor cortex of Idoya, a monkey trained to walk on a treadmill. Neu- rons in motor cortex command muscle ac- patterns recorded from an area of her motor cortextivity necessary that controls for themovements. legs were The then firing fed into a computer that translated them into a digital output. Next, the computer receiving Idoya’s brain activity was connected via the internet to a humanoid robot, known as CB (short for Computational Brain), in Kyoto, Japan. CB locomotion and could easily walk, jump, and squat.was specifically designed to mimic bipedal

SPRING 2010 | 13 ARTICLES

Currently, Erik is able to use the “speech synthesis” system to produce vowel sounds A Voice from the Future in real-time8. Erik is presented with a target sound, such as OO (as in hoot) and A (as in hen Erik Ramsey was approached hot), and instructed to attempt to move his by Dr. Phillip Kennedy, chief sci- tongue and jaw as if he were actually produc- Wentist at Neural Signals Inc1, to ing the sound. This imagined movement acti- participate in an experimental surgery and vates his speech-motor area. The system then decodes his neural activity and generates the of his eyes. sound he is attempting to make. After train- researchDuring program, the ten hehour consented surgical withprocedure, a flick ing, Erik is able to produce the target vowel a “neurotrophic electrode” was placed within Erik’s speech-motor area . The speech- The next step for Erik and the researchers motor area coordinates the 7muscles of the issounds the production with an 89% of consonants;accuracy. the ultimate jaw, tongue, and throat that are necessary goal is words. Each milestone requires the de- for the production of speech. The tip of the velopment of new speech processors and de- recording electrode was placed 5mm into coding algorithms. Erik’s cortex. The outer end was attached Erik and patients like him are living proof to an FM transmitter which could wirelessly of the emerging coalescence of man and ma- relay the activity of the region’s neurons to chine. Have we reached a new plateau in the an external decoder. evolution of humanity? Think about this the After successful implantation, the neural signals of Erik’s speech-motor area could way around town or your Nike+ to track your be precisely recorded and translated into a workoutnext time progress. you use your iPhone to find your digital signal. The Neural Signals team then reached out to Dr. Frank Guenther of Boston 1. Foer J. The Unspeakable Odyssey of the Mo- University. tionless Boy. esquire.com. October 2008. Dr. Guenther had dedicated a majority 2. National Institute of Neurological Disorders of his research career to the development and Stroke, http://www.ninds.nih.gov/ of an accurate mathematical representation disorders/lockedinsyndrome/lockedin- of the neural signals associated with speech syndrome.htm production. His expertise on the brain interactions necessary for speech production Neural Interface Systems. Annu. Rev. Neu- was of paramount importance in the cre- 3. Hatsopoulos,rosci N. Donoghue, J. The Science of ation of algorithms and software capable of 4. National Institute on Deafness and Other interpreting neural activity in Erik’s brain. Communication. 2009. 32:249–66 Disorders. http://www. “We can decode intended sound di- nidcd.nih.gov/health/hearing/coch.asp rectly from the premotor cortex [where the 5. M Kawato. Brain controlled robots. HFSP J. speech-motor area is found], rather than having to decode intended muscle activity. 6. Blakeslee S. Monkey’s Thoughts Propel Ro- This suggests that when you talk to yourself bot,2(3): a136–142. Step That 2008. May Help Humans. NY in your head, your premotor cortical neu- Times. Jan 15 2008. rons are representing the sound that would 6. Bartels J et al. Neurotrophic electrode: come out of your mouth if you were actually Method of assembly and implantation speaking the words. Furthermore, we can into human motor speech cortex. Jour- tap into this ‘inner speech signal’ and send nal of Neuroscience Methods it out of a computer speaker, thereby allowing paralyzed individuals to regain speech . 174(2):168-- capabilities,” Dr. Guenther told me in an e- chine176. 2008. Interface for Real-Time Speech Syn- mail. 7. Guentherthesis. PLoS FH, ONE. et al. A Wireless Brain-Ma

4(12): e8218. 2009.

14 | THE NERVE ARTICLES

Brain Battles: The Ethical Clash of Neuroscience and the Military

By Monika Chitre

SPRING 2010 | 15 ARTICLES

American males and ten percent of have the ability to take pills that American females experience post- would expand his working mem- traumatic stress disorder (PTSD), ory and learning while increasing and these percentages are even productivity? Would these pills be higher among high-risk popula- available equally to everyone, re- tions like war veterans2. gardless of socioeconomic status? The military has maintained a Or would those who are already discrete relationship with the mind high achievers simply propel themselves forward in a vicious feed- Dr. Jonathan Moreno, a professor forward cycle? These questions atand University brain sciences of Pennsylvania since the 1950’s. and do not even touch on the concern the author of the 2006 book Mind of where society would be headed told a story and shown a slide- Wars, talks about how his father if such drugs were on the market. Ishown 1997, about a group a series of people of events. were tested LSD for the military . Over On the other hand, a smaller effort one third of psychologists received3 has been put into the development car accident was described, but the funding from the government in of agents that can interfere with beginningIn the middle and of the ending story, portion a horrific of - memory formation. the story were emotionally neutral search in the military. During the The primary role suggested in context. One third of the group hopes of finding a use for their re for such a drug would involve pa- was given a placebo one hour be- developed interrogation tech- tients who suffer from PTSD, which fore the story was told. One third niques1950’s that and involved beyond, psychological psychologists is extremely prevalent in veterans . of the group did not receive any torture and humiliation . In the In 2002, a study was conducted9 drugs. And the last third received 21st century, the intimate3 relation- with PTSD patients involving pro- ship between the mind and brain pranalol. The patients were given blocker typically used to decrease sciences and the military was the drug in addition to talk therapy thepropranolol effects of hypertension. – a beta-adrenergic A week touched upon when the American over the course of three months. later, the subjects who were given Psychological Association refused Initially, little difference in the pa- the placebo or no drugs at all re- to comment on the torture scandal tient’s psychiatric evaluations was membered the emotional portion that took place at the Iraqi prison shown between the placebo and Abu Ghraib . - well. However, those who received Currently,3 the military has cant change occurred at the three- the– the propanolol car accident scored – lower of the on story the shown interest in the development propranalol groups, but a signifi- memory test than any of the other of a drug that would dampen con- bo recipients responded to trigger participants1. solidation of emotional memories stimulimonth follow-up. (recording 43% of of the the patient place When American talking about soldiers leave the the traumatic country for deploy- “ n e u ro s c i e n t i s t s event), while ment, they undergo none of the a certain amount of p ro p ra n a l o l mental stress that can change the way subjects re- most civilians will sponded phys- never have to en- i o l o g i c a l l y. dure. The days are Although the long and physically data was not exhausting. Many robust, it was soldiers must cope with the fact in soldiers. While most of the re- apparent that such a drug could that they have killed human beings. - Some soldiers may even have to in- on drugs that enhance cognition by fect on those suffering from PTSD4. terrogate prisoners and essentially actingsearch onsince dopaminergic the 1960’s has pathways, focused haveIn a Mind significant Wars, attenuation Moreno men ef- torture them to drawAmerica more infor- fights wars”tioned that the use of this drug mation out of them. Five percent of the most controversial. Should one could be taken a step further. In- this field has also turned out to be

16 | THE NERVE ARTICLES stead of giving the drug to dampen his Stanford prison experiment in American physicists wrote the Rus- the effects of PTSD, the drug could - sell-Einstein Manifesto to voice the be administered as a preventa- sively. He concluded that most (if concerns and consequences that tive measure. Soldiers can take the not1971 all) – researched individuals the are event capable exten of come with atomic bombs. Many drug before they go to battle or in- treating others the way the soldiers of the Manhattan Project physi- terrogate a detainee. After taking treated the prisoners at Abu Ghraib cists grew sober after the atomic the drug, they would not maintain because of the system in which an bombs they created were actually feelings of guilt or distress after individual participates6. used. They intimately understood they kill an enemy or torture a pris- “If you put good apples in a bad all of the dangerous after-effects oner . On the other end of interro- situation, you’ll get bad apples,” that these bombs would cause the 3 “soldiers taking propranolol would remember everything that happened, but those memories would have no emotional value” gation, oxytocin could also be used said Zimbardo in an attempt to ex- world if countries continued to use to increase the amount of trust the plain an individual’s capacity for these weapons extensively. Such prisoner has in the interrogator, so evil in an interview with the New awareness does not exist in the he may give up more information York Times6. neuroscience community yet, es- easily5. Importantly, propranolol However, neuroscientists fail pecially because the line between only the attenuates the emotional to realize that the products of their offensive and defensive use of neu- component of memories, not the research can and most likely will roscience is clouded . memory itself; thus, the soldiers be abused in the military. In Mind 3 taking propranolol would remem- Wars, Moreno comments that his - ber everything that happened, but neuroscientist colleagues cancel In the late 1980’s and early those memories would have no seminars on the ethics of advanc- Britain1990’s, becausethe Soviet of the Union research came they un emotional value10. es in neuroscience because only a wereder fire conducting from the onUnited dangerous States bioand- Though beta-blockers seem handful of people plan to attend . logical substances like smallpox. like a good idea in theory, the state The National Academy of Sciences3 Their work was being done behind of American interrogation tech- did not recommend that the mili- a curtain because they were using niques allows the question of eth- tary take more interest in neurosci- ics to arise. Such questions came to . In compar- tune bioterrorist techniques for the a head after the Abu Ghraib torture 7 Sovietthis research military. exclusivelyOnce the USSR to fine- fell, scandal in 2004. Once the infor- ence until May of 2009 - the secrets came out, and the re- mation was exposed to the public, ateison atomic to the physicistsbombs for of the the military, 1950’s search was stopped. In the United the American military panicked neuroscientistswho were hired havespecifically not come to cre to States, it has been decided that any understand the importance of their research on biological compounds blame. It was not clear if the scan- research to the government as must be limited so that it can only daland occurred scrambled simply to find because a source of to a quickly as they should have . be used defensively by the Ameri- few intrinsically evil individuals, Such ignorance may 3 impede can military. Unlike bioterrorism, the environment, or the people neuroscientists’ research from be- neuroscience can easily be used who create the environment itself. offensively and defensively. In the case of interrogation, psychological ing used ethically. In the 1950’s, Dr. Philip Zimbardo – famous for Einstein and a panel of high-profile SPRING 2010 | 17 ARTICLES torture and humiliation tactics are Ghraib could happen. the American military by not tor- used commonly, and it is question- Although such a drug could turing him. This article, written in able if this use is purely defensive . prove very useful for defensive tac- 2008, claims that few changes have With the advent of drugs like pro3- tics and treatment of PTSD, the mil- taken place in the military’s inter- pranalol, the results could be even itary should also use neuroscience rogation techniques despite the worse. and social sciences to create new success that Alexander brought to Abu Ghraib happened without techniques for interrogation. Mat- the mission in Iraq8. any drug intervention. The soldiers thew Alexander, an interrogator The relationship between neu- who exhibited extreme sadistic responsible for taking down Zar- roscience and military will never tendencies by physically and sexu- qawi’s forces in 2006, commented disappear. However, neuroscien- ally abusing their prisoners did not in an article to the Washington tists can take advantage of their need drugs to numb their emotions Post that he could not stand what ever-growing ties with the military because the environment and the was going on in Iraq. Alexander to ensure that their research won’t system that created the environ- goes on to explain that the reason be abused by the government. If ment easily pushed them into being he was successful in subduing Zar- neuroscientists take a more active monsters6 qawi was because he used his own role in execution of their research effects of propranalol would most brand of interrogation technique in the military, they could essen- likely make. A drug interrogation with intensified even that capitalized on cultivating trust tially change the way America worse. If soldiers cannot feel the in the detainee. One detainee told emotional horrors they create and Alexander that the reason he gave fail to feel guilty about the things Alexander information was be- fights wars. they do, scandals worse than Abu

cause he defied the stereotype of

1. LaBar, KS and Cabeza R. “Cogni- mood and pro-social behavior Iraq. The Washington Post. tive neuroscience of emo- and blunting of unwanted tional memory.” January memories. Neuroscience and - Nature Reviews: Behavorial Reviews lenceNovember of post-traumatic 30, 2008. stress Neuroscience. 9. Schlengerdisorder WE in theet al. Vietnam The preva 2. “PTSD2007: Fact 56-58. Sheet.” http://www. 6. Dreifus, Claudia. Finding. 32: Hope 760- generation: A multimethod, athealth.com/Consumer/dis- in766, Knowing 2008 the Universal Ca- multisource assessment of orders/ptsdfacts.html pacity for Evil. The New York psychiatric disorder. Journal Times. of Traumatic Stress Dana Press. 2006. - 4.3. Doskoch,Moreno, Jonathan. Peter. Can Mind Beta Wars. ing Neuroscience April 3, 2007. To Improve 10. Farah MJ et al. Neurocognitive, 5, 333- Blockers Prevent PTSD? A 7. Pincus,Soldiers’ Walter. Performance. Study Urges The Us enhancement:363, 1992. What can we First Look. Neuropsychiatry Washington Post. May 18, do and what should we do? Reviews Nature Reviews Neuroscience 5. Jongh et al. Botox for the brain: 8. Alexander, Matthew. I’m Still 5, 421-425 (2004). enhancement. 3(2), of2002. cognition, Tortured2009. by What I Saw in

18 | THE NERVE ARTICLES

Neuroscience and Religion: Using Brains to Explore the Divine

By Pinkey Shah

SPRING 2010 | 19 ARTICLES

ewish philosopher Martin Bu- tranquility from feeling personal “father-complex”. As a helpless ber said that “[M]an cannot “closeness” to the divine as well as child looks to his father for love and Japproach the divine by reach- the fear of divine retribution are protection, an individual who fears ing beyond the human: he can ap- his own fate looks to God for pro- proach it through becoming hu- and security of a higher power . tection from life’s perils. Freud ar- man. To become human is what he, Rites,reinforcements rituals, or of prayers the magnificence may allow3 gues that religion also breeds con- the individual man, has been creat- - formity in its followers through its ed for.”1 Buber’s is just one of many tion in this greater scheme and in promises of reward in exchange for voices trying to determine the ori- turnindividuals create anto influenceenvironment their condu posi- general compliance 4. This model of gin and the purposes of religion. cive towards personal growth and the Western world’s interactions Buber believes that understanding happiness. This appeal of religion between society and religion held religion is understanding oneself. offers a concrete sense of purpose true until the early seventeenth and an overarching “universal discern whether he is challenging plan”8 which provides a measure began to challenge religious doc- religiousHis vague zealotswords make by arguing it difficult that to of security in an otherwise unclear century when scientific advances their ideas cannot possibly reach world. Accordingly, many religious The Galileo Affair, perhaps beyond the scope of the human followers welcome the notions thetrine most on a significant well-known scale. clash of re - mind, or whether he is actually of “destiny” and even the “soul,” praising religion for its essential which carries their essential being in the West, began in 1610 after simplicity and attractive way of life. onward after death. These beliefs Galileoligious andpublished scientific Sidereus authorities Nun- Regardless, many theologists and can largely determine the conduct cius (Starry Messenger). The pa- philosophers agree with Buber’s of many people. The shared expe- per contained observations Galileo principal argument regarding the riences like-minded believers take made via telescope that supported - part in create an important sense the theory of a heliocentric model gious beliefs. of communal unity, and both scien- of the universe. The heliocentric reflectionTo discuss of human “religion” nature in as reli a theory was originally proposed phenomenon, one must clarify linked faith, religion, and spiritual- by Copernicus and argued that the boundaries of the term. The itytific to and both anecdotal physical and evidence emotional have the Earth and the planets revolve well-being. around the Sun while the Sun is set of beliefs shared by a common While many view religion as stationary at the universe’s center. people.broadest In definition many cases, of religionthese belief is a a system that supports happiness This idea was in clear contrast with systems seek to explain the nature and comfort even after death, the the Catholic Church and its beliefs of existence12. The large percentage institution is not without its op- concerning a geocentric universe, of Americans that report a belief ponents; after all, countless wars a theory proposed by Ptolemy in some sort of religion demon- and atrocious crimes have been and supported by the Christian strates the appeal of the varying committed in the name of God, the canon. Geocentrism argues that religions. Statistics have suggested Crusades being just one example. the Earth is the center of the uni- that as many as ninety-six percent One prominent opponent of reli- verse and all of the heavenly bod- of Americans believe in some form gion was psychoanalyst Sigmund ies revolve around the Earth. This of a god or universal spirit 8. Many Freud, who regarded religion as belief centered around the impor- religious groups share three pri- a compensatory mechanism for tance of Earth in the Bible as well mary features: a belief in an after- society’s inability to maintain or- as several scriptural passages that life, rituals and demonstrations of der and achieve productivity. His made reference to Earth’s apparent faith, and an attribution of events stance states that humans natu- - rally behave in a destructive and parent mundane causes and rami- chaotic manner when they lack fixed state. Unable to prevent the to greater. significance than the ap a governing central power. As a bespread taught of Galileo’s as a hypothesis findings, onlychurch in To some,9 religion offers emo- result, religions aim to establish officials allowed heliocentrism to- tionalfications and mental stability tran- an appropriate code of conduct. ing its spread to general populace scendent of simple human in- Freud calls human nature’s need inscientific an attempt circles, to prevent while prevent discord teractions and mentalities. Both for religion to maintain society the within the Christian community. In

20 | THE NERVE ARTICLES “an important ramification of our pattern-seeking is the inevitable search for understanding of the human existence and explanations for unsolved problems” the end though, Galileo was put on portion of the brain. Lower level claimed the search for a “God gene” brain structures include the mid- based on the apparent conserva- of heresy for his claim against the brain, cerebellum, and brain stem. tion of a parent’s religious com- teachingstrial in 1633 of and the was Catholic found Church guilty The midbrain contains the limbic mitments in his children. A child, and the Holy Bible2. system, which is evolutionarily old- for example, whose father serves Neuroscience has taken an er than the neocortex and sits be- as a pastor, will likely grow up in interest in examining man’s rela- tween the neocortex and the brain a religious setting. Now, this child tionship to religion through the stem and is responsible for much of may voluntarily, or perhaps invol- brain given its apparent role as emotion and memory 1. untarily, share the same beliefs as the “bridge between the physi- Investigations of the neurosci- his father, and the question of “na- cal and the imaginative.”1 A neu- ence of religion often logically fo- ture versus nurture,” arises10. The roscientist might say that lower cus on the limbic system due to the hypothesized “God gene” would neural networks process external strong association of religion with include inherited genetic informa- and internal stimuli and modulate emotions and emotional memory. tion that predisposes individuals to simple, instinctive behavior, while “Sacred emotions” describe genu- believe in a higher power. This hy- the higher ones, such as the frontal ine emotions associated with reli- pothesis was formulated by geneti- lobes, provide self-awareness and gious commitment and result from cist Dean Hamer who offers four the great range of thought that sep- consistent exposure to religious major pieces of evidence to sup- arates man from other animals. It is port his theory, the most relevant necessary then, to build the model as “emotional appreciation and of which is the discovery of a gene of the higher-lower functions from thankfulnesssettings. Simple for gratitude favors received.”is defined known as VMAT2. This gene codes a basic level to discuss religion’s A “sacred” form of gratitude may for a transporter of monoamine complex relationship with human result from religiously-inspired im- neurotransmitters in the brain and cognition. portance of gratitude for everyday this transporter alters critical brain To provide a brief overview of items and privileges that might be chemistry, which demonstrates it- the relevant brain structures, the otherwise taken for granted, such self ultimately in behavior. Hamer six-layers of cells associated with as food, clothing, and shelter. This called attention to the gene when sensory perception, motor com- heightened sense of gratitude may he found that it was a common ele- mand and planning, spatial reason- promote positive emotions, result- ment in the DNA of over 1000 in- ing, conscious thought, and lan- ing in increased enthusiasm and dividuals claiming to be devoutly guage is called the neocortex. This energy. From a medical perspec- religious. The data is still out on feature is evolutionarily the new- tive, the chemicals responsible for how strong a connection exists be- est and contains the most “higher- the positive feelings associated tween VMAT2 and faith, but Hamer order” functions. The frontal lobe, with gratitude may alter functions does not expect to see a direct link located right behind the forehead, as essential as pulse, blood pres- - is responsible for most abstract sure, and even muscle tone. Grati- ality; rather, he believes his work thought such as the recognition tude and many other emotions all between his findings and spiritu- of cause-effect relationships, dis- show varying kinds of widespread rological pathway connecting the tinguishing “good” from “bad”, effects on man’s autonomic ner- humanis a step intellect towards and finding investment the neu in and essential logic and reasoning vous system. 11 religion5. skills.. Other parts of the neocor- Many scientists have turned tex include the parietal lobe in the argues that genetic information toward studies of the brain in ac- superior and posterior portion of guidingPlenty religious of scientific behavior speculation exists tion rather than trying to identify the brain, the temporal lobes on and that when these genes are ex- genes that correspond to religious the sides of the brain, and the oc- pressed one tends to exhibit reli- piety, in order to examine the neu- cipital lobes in the farthest back gious behaviors10. Some have even

ral roots of religion. The field of SPRING 2010 | 21 ARTICLES brain mapping has grown rapidly above all else, a pattern-seeking have attempted to examine the as technology allows incredibly - origins of religion and its complex clear studies of brain chemistry and ish musical scales; feel uneasy lis- role in understanding the human electrical signaling. The ultimate teningentity. Weto random feel compelled noise; and to finsee condition. Ideally, neuroscientists goal is to establish a clear connec- wholesome shapes when looking hope to establish a clear biological tion between precise areas of the at partially occluded objects. But explanation for the rise and perse- verance of religion as a universal workings of human consciousness. pattern-seeking is the inevitable phenomenon throughout history. Perhapsbrain and if the how neural they center influence for “an the- searchan important for understanding ramification of the of ourhu- Looking ahead, one wonders ger” as an emotional phenomenon man existence and explanations for how religion and man will interact were to be located and changed unsolved problems. This may be in as science solves more and more so as to remain inactive, one could part responsible for the universal- of the universe that long needed prevent the emotion, serving as ity of religion in human culture and religious faith to explicate. When the ultimate anger-management in turn may have aided science’s individuals feel the need to seek program. In regards to studies of rise to predominance in the West- guidance, will they consult reli- - ern world. Science concerns itself dence gathered at the University of with experimental data and theory As neuroscience examines religion Californiareligious experience, in San Diego scientific points to evi an while religion claims authority for moregious authorities carefully, willor scientific people ones? share area in the temporal lobe, though the supramundane concepts con- not a region traditionally associ- sidered beyond the scope of man8. construct more and more? The in- ated predominantly with emotion, If an explanation for some prob- credibleFreud’s view longevity of religion of religious as a flimsy be- that produces “intense feelings of lem isn’t evident, God must be re- havior suggests a functional per- spiritual transcendence combined sponsible. Religious belief cannot manence to its existence in society with a sense of mystical presence.” and that it may very well never One neuroscientist, Michael Pers- single gene, but rather exists as a disappear from its prominent po- inger of Laurentian University, was consequencebe confined to of the some expression level of of hu a- sition in everyday life. Elucidation able to reproduce these feelings man nature, the need to explain the of the neural bases of religious be- in individuals with no reported inexplicable - lief promises only to explain why religious devotion by stimulating tions of the universe,7 man, and even people tend to be religious. Fig- the appropriate area within the religion spread,. As religionsscientific evolveexplana in uring out why and how the brain temporal lobe. Many participants their own way to continually of- makes one believe in a god does claimed feeling a greater “pres- fer support in avenues in which not demean that belief in any way; ence” and, in one such experiment, they claim science lacks author- nor does it prove that God doesn’t a participant reported “[seeing] exist. Perhaps the most pressing discovery has caused religion to question is how politicians and in front of him as this brain region ity. Freud believed that scientific the public will respond to such re- wasChrist stimulated in the strobe”. light flashing the ever-growing religious popula- search and whether scientists will The human3 brain may be, tionlose4 .much Psychology of its influence,and neuroscience despite be supported in their quests.

1. Ashbrook, James B. and Carol Rausch Albright. Hardwired into our Genes. New York: Dou- Meet: Evolution, Genes, and the Religious The Humanizing Brain: Where Religion and bleday. 211-12. Brain. Westport: Praeger. Vol. 1. 2006. Neuroscience Meet. Pilgrim Press. Cleve- 6. Heinze, Andrew R. “Jews and American Popular 10. McNamara, Patrick. Where God and Science Psychology: Reconsidering the Protestant Meet: The Neurology of Religious Experi- Paradigm of Popular Thought”. The Journal ence. Westport: Praeger. Vol. 2. 2006. land, Ohio. 1997. and the Bible. Notre Dame, IN: University of 11. McNamara, Patrick., ed. Where God and Sci- 2. Blackwell, Richard J. (1991). Galileo, Bellarmine, Notre Dame Press - ence Meet: The Psychology of Religious Ex- of American History. Vol. 88. 2001. 950-978. rience: A Study in Human Nature. New York: 7. James, William. The Varieties of Religious Expe 12. Pargament, Kenneth I. “Is Religion Nothing 3. Carter, Rita. Mapping the Mind. University of perience Westport: Praeger. Vol. 3. 2006. 4. Freud, Sigmund. The Future of an Illusion. To- 8. Long, Edward Leroy. Science and the Christian but…? Explaining Religion vs. Explaining Re- California Press. Los Angeles. 1998. Random House. 1929.

5. Hamer, Dean H. The God Gene: How Faith is ronto: Hogarth Press. Vol. 21. 1961. Faith. New York: Haddam House. 1950 ligion Away”. Psychological Inquiry. Vol. 13. 9. McNamara, Patrick. Where God and Science 2002. 239-44

22 | THE NERVE ARTICLES

Depression have you in a funk? Try the Special K Challenge.

By Aisha Sohail and Jeff Wessell

SPRING 2010 | 23 ARTICLES

Introduction step. Ultimately CREB, a transcription factor, binds to a region of DNA inside the cell and increases Brain-De- n high school, they warned us of the dangers of rived Neurotrophic Factor (BDNF). BDNF is a neuro- date rape drugs such as Rohypnol, GHB and Ket- trophin involved in promotion of neurogenesis, differ- Iamine. But current studies demonstrating the clin- entiation of new neurons and synapses, and neuronal ical uses of these once illicit drugs shed light for their growth. The BDNF hypothesis of depression is rational- has been shown to be a potent hypnotic and is used ized by hippocampal neuronal atrophy and death. inuses foreign as pharmacotheraputics. countries to treat patients Rohypnol, with orsevere roofies, in- somnia. GHB is now FDA-approved to treat patients in emotional control,1 therefore adverse neuronal with narcolepsy. But what may be most intriguing is changesThe hippocampus can account is believed for mood to disorders.play a significant Decreased role that Ketamine (aka Special K) has gone through sev- neurotrophin levels are correlated with a lack of neu- eral clinical trials that implicate it as a powerful anti- rogenesis in a given brain region. Thus, current an- depressant, working especially well in patients with treatment-resistant depression. by increasing neurotrophin levels and eventually promotingtidepressants neurogenesis. fit the BDNF Clinical hypothesis evidence of depression in support of the BDNF hypothesis has been presented by Chen Traditional Antidepressants et al using immunohistochemistry to stain for BDNF levels in post-mortem patients with mood disorders urrently there are three main classes of antide- who were either treated or untreated with antidepressant treatments that are prescribed to pa- pressants.2 Subjects with mood disorders on antide- Ctients with major depressive disorder (MDD). pressants showed increased levels of BDNF in regions MDD is a disorder characterized by overall low mood of the hippocampus compared to non-medicated indi- and can be accompanied by a lack of interest or plea- viduals with mood disorders. sure, and low self esteem. The three classes of treat- Current antidepressants do not effect a change in ments - Monoamine Oxidase Inhibitors (MAOIs), individuals immediately; patients must be taking the Tricyclics, and Selective Serotonin Reuptake Inhibi- medication for 2-4 weeks before full therapeutic effects are attained. In a study performed by Wang et were designed to increase levels of neurotransmitters availabletors (SSRIs) at synaptic – share junctions a common because final pathway;for years, “low they to male mice demonstrated an increase in dendritic levels of neurotransmitters” has been the hallmark of arborization,al, chronic fluoxetine an indicator (Prozac, of an neuronal SSRI) administration activity and depression. However, recent research shows that the proliferation. story is not that simple. 3 Individuals with depression have a decreased for the neuroplastic Only chronicchanges fluoxetine and increased produced neuronal such volume in their prefrontal cortex which is accompa- complexityresults after to 3-4 take weeks, place. suggesting This result thatcomplements it takes time the nied by neuronal atrophy. Until recently, this neuronal time necessary for the medication to enact its behav- degeneration was believed to be irreversible and un- ioral effects in humans. It is assumed that this delay treatable. New studies have proved that neurogene- period is one in which downstream effects required to sis, a process of new neuronal generation and growth, produce neuroplastic changes and neurogenesis oc- can occur in mature brains. All of the current forms cur, suggesting that therapeutic effects cannot be at- of antidepressant treatment indirectly cause neuro- tained in a short period of time. genesis by elevating synaptic neurotransmitters. For instance, SSRIs selectively block reuptake of serotonin (5HT) thus increasing the amount of 5HT in the syn- Evidence for Ketamine as a potent antidepressant apse. The elevated amounts of 5HT cause a cascade of postsynaptic activity beginning with elevating levels arate, Singh, and colleagues (2006) found a ro- of cAMP, an important intracellular second messen- bust, rapid (hours), and relatively sustained (1 ger. Elevated levels of cAMP in turn increase levels Zweek) response to a single dose of the NMDA of intracellular PKA, which in turn increases levels antagonist ketamine in patients with treatment-re- of CREB. This process is not merely a domino effect, sistant MDD; subjects infused with ketamine showed but involves various regulatory factors between each

significant improvements on a depression rating scale 24 | THE NERVE ARTICLES test compared to those infused with placebo. antidepressant actions. The Forced Swim Test (FST) is These results are remarkable considering ket- a paradigm to measure depression in which immobil- amine’s short half-life of only 2 hours. It is unlikely ity (swimming) time correlates well with depressive that ketamine’s euphoric effects were mistaken for symptoms in mice. When NBQX was administered its antidepressant properties because ketamine’s antidepressant effects last for up to 1 week. These ef- in immobility time compared to placebo; however, fects also cannot be attributed to the placebo effect, as whento mice NBQX before was FST, administered there was no immediately significant prior change to ketamine administration, it abolished the decrease in over placebo, and the effect sizes of this study were immobility time.8 Common treatments with standard various analyses showed the significance of ketamine antidepressants have also been shown to enhance post-administration. Similar results have been repli- AMPA receptor surface levels. This period of latency catedvery largein mouse at day models. 1 and4 moderate to large at day 7 necessary to achieve antidepressant9 effects with standard agents may represent the time necessary to exert direct modulatory effects on AMPA receptors. These Mechanisms of Ketamine’s antidepressant action observations suggest that ketamine’s antidepressant properties may be modulated by increased AMPA re- etamine binds to the NMDA receptor with an ceptor throughput. Not only does ketamine work rapidly, but it also Kthan other sites; blocking opiate, cholinergic, has sustained effects that can be explained by AMPA- or otheraffinity monoamine that is much receptors higher does for the not NMDA interfere site related neuroplastic changes. AMPA potentiation in with ketamine’s behavioral effects,5 suggesting that ketamine’s antidepressant action is consistent with its behavioral effects are probably mediated by via the the observation that AMPA may promote increased NMDA receptor. In addition to being a noncompetitive neuronal survival, attenuate apoptosis, and prevent NMDA antagonist, ketamine also produces a presyn- synaptic deterioration produced by deafferenta- aptic release of glutamate, which is tion10,11,12. AMPA receptor activation apparently essential to its antidepres- “results from increases BDNF expression. In one sant action.6 The rapid antidepressant study, AMPA stimulation in hippocam13 - effects of ketamine could be due to an ketamine pal slice cultures caused the selective increase in glutamatergic through- activation of MAPK through the up- put due to the increase in ionotropic promise to stream activator MAPK kinase (MEK). glutamate AMPA receptors relative to Inhibition of either component of the ionotropic glutamate NMDA recep- revolutionize AMPA-MAPK pathway increased cel- tors. NMDA receptors are permeable lular damage due to serum depriva- to Ca2+ ions and Na+/K+ ions, where- depression tion, suggesting that this pathway as AMPA receptors are permeable to facilitates compensatory signals in only Na+/K+ ions. NMDA receptors treatment” response to injury. Likewise, AMPA have a magnesium block mechanism potentiation with ampakine CX516 such that they are not activated until membrane has enhanced MAPK activation and reduced synaptic and been depolarized via AMPA current neuronal degeneration resulting from excitotoxic epi- If NMDA receptors are blocked by Ketamine, glu- sodes.14 Moreover, AMPA receptor activation has been tamate throughput is forced to prefer AMPA receptors shown to increase BDNF mRNA in a manner depen- over NMDA receptors. In other words, glutamate re- dent upon MAPK signaling; and BDNF-induced cell leased from a synapse will activate mostly AMPA re- survival has been shown to require the MAPK signal- - ing pathway in neuronal cultures deprived of survival tion that post-mortem studies of depressed patients factors15.16. ceptors. This finding is consistent with the observa Traditional antidepressants and glutamatergic antidepressants are thought to act via a similar path- responsibleconfirm decreased for decreasing levels ofNMDA metabotropic receptor activity. glutamate way in the brain. The traditional antidepressants receptorsIn addition, mGluR2 studies and mGluR3, with an which AMPA are receptor primarily an7 - change levels of synaptic neurotransmitters to ef- tagonist, NBQX, also demonstrate the need for in- fect a change in intracellular cAMP, PKA, CREB, and creased AMPA receptor throughput for ketamine’s ultimately BDNF. Glutamatergic antidepressants en-

SPRING 2010 | 25 ARTICLES hance glutamatergic throughput by upregulating the number of AMPA receptors with respect to NMDA re- 8. Maeng S, Zarate CA, Jr, Du J. Cellular mechanisms ceptors, increasing the MAPK intracellular signaling underlyingtivation. Nat. the Neurosci. antidepressant 2: 44 – 49. effects32. 1999. of ket- pathway. The effect is that neuronal apoptosis is at- - tenuated and levels of BDNF are increased, promoting lisoxazole-4-propionic acid receptors. Biol Psy- overall neuronal health. amine: role of alpha-amino-3-hydroxy-5-methy Results from ketamine, and soon perhaps other non-traditional antidepression drugs, promise to Signalingchiatry. 2007. in Mood Disorders. In Finkel T and Gut- revolutionize depression treatment. Perhaps in the 9. Dukind J, GouldG, editors. TD, ManjiSignal HKTransduction (2004): Neurotrophic and Human near future, ketamine therapy may defeat the chronic Health. New York, NY: John Wiley and Sons. blues that is depression. Whether or not depression 10. Bambrick, L. L., P. J. Yarowsky, and B. K. Krueger. is a phenomenon that should be eradicated is another - question. vival factor: An inherited defect in the trisomy 16 1995. Glutamate as a hippocampal neuron sur 11. Limatola, C., M. T. Ciotti, D. Mercanti, F. Vacca, D. 1. Duman RS. Depression: a case of neuronal life and Ragozzino,mouse. Proc. A. Natl. Giovannelli, Acad. Sci. A.USA Santoni, 92: 9692–9696. F. Eusebi, death? Biol Psychiatry 2004; 56: 140-145. and R. Mildei. The chemokine growth-related 2. Chen B, Dowlatshahi D, MacQueen GM, Wang JF, gene product beta protects rat cerebellar gran- Young LT. Increased hippocampal BDNF immuno- ule cells from apoptotic cell death through alpha- reactivity in subjects treated with antidepressant - medication. Biol Psychiatry 2001; 50: 260-265. - 12. McKinney,amino-3-hydroxy-5-methyl-4-isoxazolepropio R. A., M. Capogna, R. Durr, B. H. Gahwil- tion and synaptic plasticity of adult-born hippo- er,natereceptors. and S. M. Thompson. PNAS. 97: Miniature 6197– 6201. synaptic 2000. events 3. Wangcampal JJ et granule al. Chronic cells. fluoxetine The Journal stimulates of Neuroscience matura maintain dendritic spines via AMPA receptor acti-

4. Maeng S, Zarate CA, Jr, Du J. Cellular mechanisms - underlying28: 1374-1378. the 2008. antidepressant effects of ket- vation. Nat. Neurosci. 2: 44 – 49. 32. 1999. - 13. Legutkoa novel B, AMPA Li X, Skolnick receptor P. potentiator. Regulation of Neurophar BDNF ex- lisoxazole-4-propionic acid receptors. Biol Psy- pression in primary neuron culture by LY392098, amine: role of alpha-amino-3-hydroxy-5-methy 14. Bahr BA, Bendiske J, Brown QB, Munirathinam S, 5. Hustveit O, Maurset A, Oye I. Interaction of the chi- Cabamacology E, Rudin 40:1019 M, et –1027. al (2002): 2001. Survival signaling ralchiatry. forms 2007. of ketamine with opioid, phencyclidine, and selective neuroprotection through glutama- sigma and muscarinic receptors. Pharmacol Toxi- 15. Hayashi, T., H. Umemori, M. Mishina, and T. Yama- 6. Maeng S, Zarate CA, Jr, Du J. Cellular mechanisms tergic transmission. Exp Neurol 174:37– 47. underlyingcol.; 77:355-359. the antidepressant1995. effects of ket- and signals through the protein tyrosine kinase - moto. 1999. The AMPA receptor interacts with lisoxazole-4-propionic acid receptors. Biol Psy- 16. Lauterborn, J. C., G. Lynch, P. Vanderklish, A. Arai, amine: role of alpha-amino-3-hydroxy-5-methy andLyn. C.Nature M. Gall.. 397: Positive 72–76. modulation of AMPA receptors increases neurotrophin expression by andchiatry. S. M. 2007. Thompson. Miniature synaptic events hippocampal and cortical neurons. J. Neurosci. 7. McKinney,maintain R. dendritic A., M. Capogna, spines viaR. Durr, AMPA B. receptorH. Gahwiler, ac-

20: 8 –21. 2000.

26 | THE NERVE REVIEWS

Recognition Memory An Analysis of Two Prominent Theories

By Danielle Miller

SPRING 2010 | 27 REVIEWS

INTRODUCTION

ecognition memory is a particular aspect of differently. According to the proposal, both are housed memory that provides the ability for an indi- within the medial temporal lobe (MTL) structures but vidual to identify a previously encountered recollection is dependent on the hippocampus, and fa- R 1 stimulus. There are two prominent theories as to miliarity relies on the perirhinal cortex . how recognition memory operates. One theory proposes that there are two distinct processes involved in the recognition of a stimulus, called recollection Evidence from Recognition Tests and familiarity1. Another theory, however, denies the independence of the two processes within recogni- One way in which recollection and familiar- tion memory. In this theory, familiarity is primarily ity differ is in the application and availability of the thought of as a weak memory, whereas recollection is processes. Familiarity typically becomes available considered to be a type of strong memory2. This paper aims to take a closer look at both theo- process that is engaged in recognizing a stimulus1. ries by exploring evidence that supports each one. Subjectsmore quickly do very than well recollection in discriminating and is usually between the firsttwo A deeper evaluation of the dual process model and items to determine which one was previously encoun- single process model will be made in order to explore tered and which one was not. Yet, when subjects need all aspects of recognition memory. The paper will to remember much more details in order to discrimi- nate between the two objects (i.e the spatial associa- understanding all aspects that are currently proposed tions with the objects) performance is reduced, sug- aboutprovide recognition the reader memory. with a background sufficient for gesting that familiarity is engaged and used effectively in making discriminations between encountered and non encountered items, but recollection must be en- RECOLLECTION AND FAMILIARITY AS TWO DIS- gaged for further processing of differences . TINCT PROCESSES The amount of time allotted to an individual3 who is recognizing a stimulus affects performance. In one n one theory of recognition memory, recollection experiment, participants were advised to reject highly and familiarity are seen as independent processes. IIn this particular theory, recollection is the process familiar items but then begin to reject as time goes on.familiar This illustratesitems4. Participants the distinction at first between accept familiaritythe highly familiarity involves a sense of the experience but lacks details.that recovers Recollection specific and details familiarity about an can event, be displayed whereas familiarity, which is a rapid, readily available process as two separate processes with evidence from recog- usedand recollection. for fast responses At first and recognition then switches memory to recollecemploys- nition tests, receiver operating characteristics (ROC) tion, which is a much slower process that allows for curves, amnesic patients, and fMRI studies. Data sup- correct responses when given longer time1. Thus, at ports that recollection and familiarity occur in two separate areas of the brain and that their usage and the items, which was an incorrect response; recollec- function are different; depending on the task at hand, tionfirst, was those employed participants only used after familiarity familiarity, and allowing accepted for each brain region may be recruited or manipulated a correct response of rejected items.

28 | THE NERVE REVIEWS

Evidence from ROC While ROC data support the idea that familiarity and recollection are two independent processes with- More experimental results that support the dual in recognition memory, brain studies of humans and process theory is the data collected from ROC curves. animals also provide further insight to this theory. In - these studies, the region that each process is thought sponses and then relating the proportion of correct to involve is investigated more closely. recognitionsROC’s involve (the collecting hit rate) recognitionto the proportion confidence of incor re- rect recognitions (the false alarm rate)5. The shape of ROC can thereby support the existence of two pro- Evidence from Brain Studies responses, a measurement of recollection can be ob- In some cases, damage to the MTL structures can tainedcesses. by Since the asymmetry recollection of supportsthe ROC curve; high-confidence in contrast, cause amnesia. Amnesiacs have severely impaired ep- familiarity can be measured by the degree of curvilin- isodic memory, which is the type of memory function earity5. In animal models, ROC’s can be manipulated life. The MTL includes the hippocampal formation, en- to show the affects that lesions have on recognition torhinal,that relates perirhinal, to the specific and parahippocampal events in an individual’s cortices8. memory. In normal rats, the ROC curve has both an Bolstering the theory that familiarity and recollection asymmetrical and curvilinear aspect to it, represent- are two separate processes, these studies show that ing the fact that the rats use both recollection and recollection is supported by the hippocampus, where- familiarity in recognition. In rats with hippocampal as familiarity is heavily dependent on the perirhinal damage, there is a loss of the asymmetry, which sug- cortex . Therefore, according to this theory, amnesiacs gests that recollection (represented by asymmetry) is with concentrated9 damage in the hippocampal area supported by the hippocampus. Since the curvilinear should show impaired recollection whereas familiar- portion of the curve is still intact, familiarity appears ity should be relatively intact. not to be affected by hippocampal damage6. It is evi- Amnesic patients with extensive MTL damage - loss of memory strength, because an increase of delay age to the familiarity process10. Since both processes (whichdent that causes this deficita weakened in recollection memory) isleads not todue a lossto the of arewill not have equally higher impaired deficits in following recollection such with damage, less damit ap- curvilinearity and retention of recollection1. The dual process model but not the single pro- located in the MTL. Instead, recollection is dependent cess model can readily explain the data from another onpears the that hippocampus recognition and is not familiarity a unified on memory the perirhinal system study. In this experiment rats were trained using the cortex, which is in the surrounding parahipppocam- ROC paradigm. Rats that had hippocampal lesions pal region . This supports the idea that recognition produced an ROC curve without asymmetry but main- consists of9 two processes that are heavily implicated tained the curvilinear aspect of it . This means that in two different areas of the brain. rats with lesions to the hippocampus7 lost recollection A case study of patient YR, who had selective bi- processes but kept the process of familiarity. In con- lateral damage to the hippocampus, demonstrates trast, control subjects that had an intact hippocampus impairment of recollection with intact familiarity11. primarily relied on recollection, producing a more Patient YR was impaired at association memory for asymmetrical curve . This reliance on recollection for different items, a task mediated by the hippocampus, normal rats was most7 likely due to the fact that the but had intact familiarity recognition between items11. rats were exposed to the same odors and mediums for Because this study shows selective hippocampal dam- multiple pairings, providing a more concrete associa- age resulting in impaired recollection with intact fa- tion with the stimulus. Thus this study supports the miliarity, the dual process model is supported. dual process model; it effectively shows that recollec- Another patient KN, who also had damage to the tion and familiarity are dissociable and differentially hippocampal region with sparing of the perirhinal affected by hippocampal damage . The maintenance cortex, showed impaired recollection and preserved of familiarity in rats with hippocampal7 lesions sug- familiarity12. Experimenters used the remember/ gests that the hippocampus is not equally involved know procedure, which requires an individual to de- in recollection and familiarity as the single process model proposes. if it is known that the item appeared but there are no clare if an item can be recalled with specific details or

SPRING 2010 | 29 REVIEWS details surrounding the event , as well as ROC analy- Evidence from fMRI Studies 13 also supports the dissociation between recollection Some studies have focused on recording the acti- andsis, both familiarity. of which suggest a deficit in recollection. This vation of certain brain areas while a participant is do- There have also been studies where the perirhinal ing a task using functional MRI. In recognition memo- cortex has appeared necessary for normal familiarity ry tasks, individuals use tasks that employ both recall function. Patient NB, who received a surgical resection and familiarity. One particular task that is commonly of the left anterior temporal lobe structures including used is the remember/ know task. the perirhinal cortex but sparing the hippocampus, Using fMRI during the remember/ know task, experimenters examine the brain regions that are activated during the “separation of location of encoding and retrieval of remembered or known the processes implicates items. Across different techniques, hippocampal the dual process model as activation is higher for those items that are re- a key player in recognition” membered (recollected) than those that are fa- showed impairment on familiarity tasks but normal miliar1. Only recollected responses produced activity function on recollection tasks14. NB was tested using in the hippocampus noticeably different than not re- remember/ know, which is a procedure that requires membered items; familiar responses, however, did not subjects to claim whether they remember or know - (the word feels familiar) a word from a list that was tive to unfamiliar items . There was, however, encod- previously presented, and ROC paradigms. This sup- ingproduce activity significant in the anterioractivity19 in parahippocampal the hippocampus gyrus, rela ports the idea that the perirhinal cortex is implicated primarily the perirhinal cortex, for familiar items20. in the familiarity process. This evidence supports the theory that recollection and familiarity are two sepa- familiar items; recollected items did not produce ac- rate processes occurring in different regions of the tivityNo other in this MTL region. structure was significantly activated for MTL structures. In hypoxic patients, in whom the hippocampus is largely damaged due to a lack of oxygen, there are RECOGNITION AS A SINGLE PROCESS - miliarity15. In some studies recollection was severely nother theory proposes that recognition mem- damagedsome differences in hypoxic in patients deficits and between familiarity recall was and nor fa- ory is composed of a single process, therefore mal16. Yet, some studies fail to report such a severe Adenying the independence of the two processes within recognition memory. It is thought that as selective to one 17process may be due to peripheral there are not two separate and distinct processes, damagedifference in inthe deficit surrounding. The reason areas thatof the the hippocampus, deficit is not but rather that recollection and familiarity support which could then affect familiarity. strong and weak memories respectively2. This theory In animal studies, in which the lesions can be claims that there is not a difference in location be- more selective, there has been a correlation between tween the elements of recognition memory, but rather selective hippocampus damage and impaired recall. that recollection and familiarity occur both in the hip- Using ROC analysis, rats with hippocampal damage pocampus and in the perirhinal cortex collectively2. had impaired recollection and, in some cases, en- Evidence that supports this theory includes amnesia hanced familiarity18. Normal rats used recollection for studies, ROC, fMRI studies, and single cell recording. tasks whereas rats with hippocampal lesions relied These studies point to a uniform process for recogni- on familiarity. This provides more concrete evidence tion memory that only differs in the strength of the that selective damage to the hippocampus does cause memory. a deficit in recollection while sparing familiarity. 30 | THE NERVE REVIEWS

Evidence from Amnesic Patients hippocampal lesions. It must be recognized, however, that this requires an assumption that the task is test- A study of 56 hypoxic patients strongly suggests 2. that recognition memory is a singular process. In this In an fMRI study recording brain activation of particular study, in which the lesions appeared limit- MTLing recollection structures and in individuals,familiarity specifically activity in the right ed to the hippocampus, 55 of the patients had impair- perirhinal cortex and bilateral hippocampus in- ments in both recall and familiarity21. Hippocampal creased as a function of increased memory strength, damage that results in a loss of both recall and famil- regardless if the memory was recollected or familiar25. iarity suggests that the two processes are in fact not When source memory success, or successful recollec- separate and occur in the same location of the brain. tion, was held constant, activity in the hippocampus Another study which involved six patients who was still noted, suggesting that hippocampal activa- had bilateral hippocampal damage resulted in an tion occurred without recollection and was reliant equal impairment of recall and familiarity22. This pro- more on familiarity based memories25. This supports vides evidence against separate processes, as it ap- the theory that familiarity and recollection do not ex- pears that both are located in the same areas of the ist as separate processes but overlap in similar brain brain. The single process theory can be supported regions. It also suggests that MTL activity occurs as further with ROC curves that outline the memory sys- a result of memory strength and is not based on the tem’s usage of strong and weak memories. type of memory encoded. However, the study only examined recollection of the originating source of the information. Therefore, it may very well be likely Evidence from ROC that other types of recollection are separate processes from familiarity. Since the researchers only examined ROC curves can be interpreted as a representa- recollection in the form of source memory, it cannot - be claimed that all types of recollection overlap with familiarity in brain regions. thetion certainty of memory of having strength. previously ROC involvement viewed a particular of confi itemdence2 levels reflects the strength of the memory, or . This im- Evidence from Single Cell Recordings plies .that Asymmetrical the ROC curves curves without reflect asymmetrystrong 23 memories (only curvilinearity)and curvilinearity produced reflects byweak brains memories with hippocam- The recordings of single cell neurons appear to be much better for detecting the differences in memory memory as opposed to an entirely different process. strength. Hippocampal neurons of epileptic patients pal lesionsAccording reflect to the an individual’ssingle process reliance theory, on stronger a weak were recorded while the patients were asked to make memories result in a higher curve than weaker memo- discriminations between old and new items. It was ries. In one study, once the overall memory strength was similar in patients with lesions in the hippocampus to the control, the graph of the ROC was asymmet- observed that certain neurons fired when new items ric (strong memory) and matched the control ROC24. were presented and other neurons fired when old This indicates that even in the absence of the hippo- itemitems failed, were; suggestingthose that firedthat tothe the hippocampus old items increased was in- campus, the curves can obtain asymmetry, suggesting volvedfiring even in item when recognition the recollection even when of the recollection location of was the that recollection is a type of strong memory. absent26. Hippocampal activation during item recognition suggests a single process occurring, which involves both recollection and familiarity. However, the Evidence from fMRI studies point to a different conclusion; these particular neu- While ROC curves are helpful in the analysis of neurons firing in the absence of recollection may also- memory strength in recognition memory, fMRI’s are 1. rons seem to fire for all remembered items, and there functioning during a particular task. In the remem- memoryfore, may is not occurring be encoding as one for process. any specific The perirhinalmemory ber/knowuseful in determiningtask, studies have specific determined brain areas that recollec that are- cortexThus, the also firing shows of nothese distinction neurons inmay processes. not mean In that re- tion is impaired more than familiarity in patients with cordings of the perirhinal cortex, there appears to be

SPRING 2010 | 31 REVIEWS no difference in the encodings of items that will later 11. Mayes AR, et al. Associative recognition in a patient with selective be strongly remembered as opposed to those that are hippocampal lesions and relatively normal item recognition. Hip- weakly remembered2. 12. Aggleton JP, Vann SD, Denby C, Dix S, Mayes AR, Roberts N, Yoneli- pocampus. 14(6): 763-84. 2004. CONCLUSIONS nas AP. Sparing of the familiarity component of recognition memory in a patient with hippocampal pathology. Neuropsychologia ecognition memory is described by two major theories: one pertaining to a dual process that 43(12):1810-23. 2005. involves a strong dissociation between recol- R 13. Skinner EI, Grady CL, Fernandes Ma.. Reactivation of context-specific lection and familiarity and another that involves a 14. Bowles B, Crupi C, Mirsattari SM, Pigott SE, Parrent AG, Pruessner JC, brain regions during retrieval. Neuropsychologia. IN PRESS. 2009 single process that describes a distinction between Yonelinas AP, Kohler S. Impaired familiarity with preserved recol- strong and weak memories. The dual process theory lection after anterior temporal-lobe resection that spares the accounts for the data obtained from both human and animal studies, whereas the single process model can- hippocampus. PNAS. 104(41):16382-87. 2007. not fully explain some data obtained from studies. associative recognition relative to item recognition in global am- 15. Giovanello KS, Verfaellie M, Keane MM. Disproportionate deficit in - cesses (recollection) within recognition memory. Spe- 16. Yonelinas AP, Quamme JR, Widaman KF, Kroll NEA, Sauve MJ, Knight nesia. Cogn. Affect. Behav. Neurosci. 3:186-94. 2003. Selective hippocampal lesions affect specific pro RT. Mild hypoxia disrupts recollection, not familiarity. Cogn. Affect. two processes, which are localized in different areas ofcific the losses brain. of Separationcertain processes of location suggest of thethat processesthere are - Behav. Neurosci. 4:393-400. 2004. implicates the dual process model as a key player in tion and familiarity in dense hippocampal amnesia: a case study. 17. Cipolotti L, Bird C, Good T, Macmanus D, Rudge P, Shallice T. Recollec recognition memory. 18. Eichenbaum H, Fortin N, Sauvage M, Robitsek RJ, Farovik A.. An ani- Neuropsychologia. 44(3):489-506. 2006. 1. Eichenbaum H, Yonelinas A.P, Ranganath C.. The medial temporal lobe mal model of amnesia that uses receiver operating characteristics

2. Squire LR, Wixted JT, Clarck, RE.. Recognition memory and the medial in recognition memory. Neuropsychologia. IN PRESS. and recognition memory. Annu. Rev. Neurosci. 30:123-52. 2007. (ROC) analysis to distinguish recollection from familiarity deficits

Remembering episodes: a selective role for the hippocampus dur- temporal lobe: a new perspective. Nat. Rev. Neurosci. 8(11) 872- 19. Eldridge LL, Knowlton BJ, Furmanski CS, Bookheimer SY, Engel SA. 83. 2007. item versus spatial position information. J. Exp. Psychol. Learn 20. Ranganath C, Yonelinas AP, Cohen MX, Dy CJ, Tom SM, D’Esposito M. 3. Gronlund SD, Edwards MB, Ohrt DD.. Comparison of the retrieval of ing retrieval. Nature Neuroscience 3(11):1149-53. 2000. Dissociable correlates of recollection and familiarity within the 4. Gronlund SD, Ratcliff R.. Time course of item and associative informa- Mem. Cogn. 235:1261-74. 1997. tion: implications for global memory models. J. Exp. Psychol. Learn 21. Wixted JT, Squire LR. Recall and recognition are equally impaired in medial temporal lobes. Neuropsychologia 42(1): 2-13. 2003. patients with selective hippocampal damage. Cogn. Affect Behav. 5. Yonelinas AP.. Receiver-operating characteristics in recognition mem- Neurosci. 4:58-66. 2004. Mem. Cogn. 155:846-58. 1989. ory: evidence for a dual-process model. J. Exp. Psychol. Learn Mem. 22. Manns JR, Hopkins RO, Reed JM, Kitchener EQ, Squire LR. Recogni-

6. Fortin NJ, Wright SP, Eichenbaum H. Recollection-like memory re- Cogn. 20(6):1341-54. 1994. tion memory and the human hippocampus. Neuron. 37:171-180. - 2003. ating characteristic in recognition memory. J. Exp. Psychol. Learn trieval in rats is dependent on the hippocampus. Nature 431:188- 23. Glanzer M, Kim K, Hilford A, Adams JK. Slope of the receiver-oper - 191. 2004. ognition memory: opposite effects of hippocampal damage on rec- 24. Wais PE, Wixted JT, Hopkins RO, Squire LR. The hippocampus sup- 7. Sauvage MM, Fortin NJ, Owens CB, Yonelinas AP, Eichenbaum H. Rec Mem. Cogn. 252:500-13. 1999. ollection and familiarity. Nature. 1:16-18. 2008. ports both the recollection and the familiarity components of rec- 8. Preston AR, Gabrieli JDE. Different functions for different medial tem- 25. Kirwan CB, Wixted JT, Squire LR. Activity in the medial temporal lobe ognition memory. Neuron. 49(3):459-66. 2006. predicts memory strength, whereas activity in the prefrontal cor- poral lobe structures? Learn. Mem. 9:215-17. 2002. the perirhinal cortex and hippocampus? Nat. Rev. Neurosci. 2:51- tex predicts recollection. J Neurosci. 28(42):10541-548. 2008. 9. Brown MW, Aggleton JP. Recognition memory: What are the roles of 61. 2001. 26. Rutishauser U, Mamelak AN, Schuman EN.. Single-trial learning of 10. Stark CE, Squire LR. Recognition memory and familiarity judgments novel stimuli by individual neurons of the human hippocampus- in severe amnesia: no evidence for a contribution of repetition

amygdala complex. Neuron. 49:805-13. 2006. priming. Behav. Neurosci. 114(3):459-67. 2000.

32 | THE NERVE The Neuroanatomy of Homosexuality

By Claire Bryson REVIEWS

he anterior hypothalamus of the human brain pressed properly, causes testicles to develop, while is sexually dimorphic and has been investigated in absence of its function, ovaries will develop.2 After Tfor dimorphism with regards to sexual orienta- - tion through examination of postmortem brains of ho- cretions direct the organization of the rest of the body, mosexual males, heterosexual males, and heterosex- includingthis stage, the the brain, testes which and their also specificdevelops hormonal either mas se- ual females. Studies examining the interstitial nuclei culine or feminine traits. As seen from the process, of the anterior hypothalamus and the anterior com- humans begin as “female,” so to speak, and develop misure4 have shown correlation3 to sexually-orientated the male characteristics. Thus, hormonal secretions dimorphism. Through further research, a relationship are both important and delicate in male development, between the development of these sexually dimorphic depending largely on the androgenic, or masculin- regions and steroid hormone levels has been estab- izing, steroid hormone testosterone.2 This molecule lished. Though neither consequence nor causation functions as a “pro-hormone” to masculinize the brain during development.2 Studies examine the occurence its implications are suggestive of the early differentia- of this neuronal patterning according to gender and tioncan beof humandefinitively sexual demonstrated orientation and from its this origins research, span- its prevalence in homosexual men in attempts to de- ning neurobiology and endocrinology. termine whether sexually dimorphic areas of the Sexual orientation has long been a topic of con- brain are affected by, or even correlate to, the sexual orientation of males. In a key article, Simon LeVay presented the differ- examinedtroversy and the debate biology in ofsocial, this political, phenomenon. and scientific Most of ence in hypothalamic structure between heterosexual thiscircles. research In the haslast focusedtwenty years, on the scientific anterior efforts hypothala have- and homosexual men . Cited by a majority of articles mus and the preoptic area (POA) within the larger 3 - hypothalamic region. Historically, these areas have tory and led to extensive additional research. LeVay shown a relationship to in the field, LeVay’s hypothesisargued and that study the was hypothala exposi- male sexual behavior.1 “there is little doubt that mus held the the key to More recent research has sexual orientation. He cited elucidated differences in these areas of the brain are that, in male monkeys, le- the anterior hypothalamus sexually dimorphic; however, sions in this region impair and its endocrinological heterosexual behavior functions between hetero- debate still exists regarding without eliminating sexual sexual males and homosex- drive . LeVay investigated ual males. In the current lit- how these differences affect 3 erature, there is little doubt of neurons, the interstitial that these areas of the brain sexual orientation” nucleitwo specific of the small anterior groups hy- are sexually dimorphic; however, debate still exists regarding how these differences affect sexual orientation. nuclei would exhibit size dimorphismpothalamus with 2 and sexual 3 (INAH ori- To understand male sexual orientation as a phe- entation,2 and 3). notHe justhypothesized with gender that as one had orbeen both previously of these nomenon, familiarity with the development of male examined . Because INAH had already been observed fetuses must be discussed. Gender is determined by to be more3 than twice as large in heterosexual men the activity of a lone pair of sex chromosomes passed than women, LeVay extrapolated that INAH 2 and/ on by the parents. Initially, the gonads are indistinct and can develop as either testes or ovaries. In males, towards women (heterosexual men and homosexual the testis determination factor (TDF) gene, when ex- women)or 3 would and be small large inin individualsindividuals sexuallysexually oriented

34 | THE NERVE REVIEWS towards men (heterosexual women and homosexual men). Because brains of homosexual women were not - available, only the brains of homosexual men were ex- erosexualhomosexual women men washad an 36% AC larger28.4% than greater heterosexual than het- amined. And although most of the homosexual male erosexualmen and 5.9% males greater4. than heterosexual women. het samples were from post mortem AIDS patients, this had no effect on the results; comparison of hetero- unknown, but it shows clear sexual dimorphism and sexual men who died of AIDS with homosexual men experimentallyThe functional determined significance sexual-orientation of the size of the dimor AC is- nuclei. Furthermore, the variance of subjects by age examined these differences in humans, but studies on waswho eliminateddied of AIDS by still age-matching showed the study difference groups. in INAH3 animalsphism, similar have shown to the that INAH they 3 area.arise Littlein the researchperinatal has or- ganism2 the hypothesized sexually-oriented dimorphism . In AC correlation to sexual orientation, the understand- accordanceLeVay’s with research his hypothesis,found that INAH the volume 3 did exhibit of 3 the ing that .no Given single the brain results structure regarding correlates both INAH to sexual 3 and nucleus was more than twice as large in heterosexual orientation carries weight. The apparent intercon- - nected nature of varying brain regions suggests that cant result . Given the double blind-procedure, no bias factors operating early in development differentiate couldmales exist,as in3 homosexual and no statistical males, differencea statistically was signifi found on the basis of gender and sexual orientation within when comparing brains of heterosexual males who sexually dimorphic structures and brain function in died from AIDS to other brains of heterosexual males. a cumulative manner2 - Ultimately, homosexual men and heterosexual women endocrinologic approach. Exploring alongside which traditional specific neuro fac- of heterosexual men, supporting LeVay’s arguments biologicaltors influence studies. which kind of development requires an both had smaller volumes. of INAH 3 compared to that The endocrinologic approach examines the role The applications of3 this discovery are limited by of sex hormones in sexual orientation. In mammalian theon the inability nature toof determineINAH 3 causation. Because of the models, like in humans, an androgen masculinizes the postmortem nature of the experiment, it was impossi- developing genitalia and then the brain as well1. The ble to gain detailed insight into all subjects’ sexualities. question is whether or not these fetal androgens alone This limits the ability to show correlation between also ‘masculinize’ the brain or if there are further key brain structure and the diversity of sexual behavior. players in development such as neurotransmitters or Additonally, “exceptions” to the rules do exist; homo- neurodevelopmental factors. to heterosexual males were observed, suggesting that turn out homosexual as a result of lower-than-normal sexual males with INAH 3 regions sized appropriately fetalConflicting androgen, fromstudies higher-than-normal have suggested that levels, boys or may for reasons having nothing to do with androgens, with impedeINAH 3 the alone ability does to identify not necessarily the dimorphism affect sexuality. of INAH no study appearing more conclusive than the rest1. Clearly, there are further unidentified variables which- However, through studies of sheep, Roselli et al.5 have gardless, sexual orientation has a demonstrated bio- some evidence that male-oriented rams receive lower- logical3 as cause correlation or consequence to some extent,of sexual which orientation. necessitates Re than-normal androgen stimulation of the brain. Since further in-depth research. testing in rats has suggested that estrogen may affect Laura S. Allen and Roger A. Gorski studied sexual the volume of the ovine sexually dimorphic nucleus dimorphism in the anterior commisure (AC) of the (oSDN), this effect was then studied in rams to explain - the minority of males which display a preference for phic but not directly related to reproductive function4. mating with other males over females5. It was found Theirhuman study, brain, similar a fiber to bundleLeVay’s, that examined is sexually volumetric dimor that male-oriented rams did in fact have a smaller nu- differences between homosexual males and hetero- cleus in this brain region than female-oriented rams5. sexuals of both genders and used age-matching and These observations lend strong support to the im- double-blind procedures to elminate variance and bias4. The AC in homosexual men was found to be sexual orientation in males5. The enzyme aromatase convertsportance androgensof the anterior like hypothalamus’s testosterone into influence estrogens on and 18% larger than that of heterosexual women4. and plays a crucial role in the masculine development When34% larger adjusted on average for individual than that brain of heterosexual mass, the AC men of of the rat SDN-POA, masculinizing the rat brain and

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“the apparent interconnected nature of varying brain regions suggests that factors operating early in development differentiate on the basis of gender and sexual orientation within sexually dimorphic structures and brain function in a cumulative manner” leading to masculine behavior1. These results show a that has not been as affected by AIDS. A longitudinal relationship between steroid hormones, oSDN mor- study would be required to track homosexual women phology, and the resulting sexual orientation in rams. to ultimately allow for post mortem analysis on the With such progress using with rams as a mammalian sexually dimorphic nucleus areas of the hypothala- model, researchers may be able to draw implications mus. Longitudinal studies could also examine varying about human sexual orientation. levels of steroid hormones throughout development. Before being applied to humans, research must There has been speculation that homosexual men in determine more precisely how steroid hormones and general have a lower age at puberty, perhaps due to the related biology affect the size of the oSDN. Does variations in hormone levels6. In general, future re- the level of aromatase activity determine oSDN size, search could also focus on the relationship between or does oSDN size determine the level of aromatase? steroid hormones and development of the SDN in the If aromatase effects in the POA are responsible for the hypothalamus. Related topics of study remain largely size of the oSDN and the size of the oSDN is in turn unexplored in humans, such as the preliminary stud- responsible for sexual preference, then interfering ies of pheromone sensitivity based on individual’s with hypothalamic aromatase in developing rams may sexual orientation . reliably shrink the oSDN and produce male-oriented The question of7 human bisexuality remains. What rams5. This demonstration would be suggestive of a biological evidence could be shown in such an individ- causal relationship between steroid action, hypotha- ual? It could prove enlightening to thoroughly exam- lamic morphology, and sexual preference in sheep5. By extension, this demonstration could also suggest that from both homosexual and heterosexual individuals. steroid hormones affect the human hypothalamus to Still,ine the correlating brain of a neurobiological bisexual person and and endocrinological find differences dimorphism cannot argue causation without further effect would further emphasize the critical biology of experimentation in animal models. Additionally, sepa- sexualinfluence orientation sexual orientation. and eliminate Studies the investigating social argument this of “choosing” one’s sexuality. gestation from purely genetic ones may not be possible.rating Therefore, environmental the interaction influences and like communication factors during done before anything truly conclusive can be reached. FirstThere of all, isthese still studiessignificant lack research homosexual that femaleneeds to sub be- be critical to future success. A concerted effort be- jects. Due to the prevalence of AIDS among the homo- tweenbetween disparate various methodologies fields of research of investigation on this topic maywill sexual male community, records of sexual orientation provide the key for advances in the years to come, and, allow for postmortem analysis of their brains. This is regardless of their results, the research itself will no not the case for homosexual women, as a population doubt have profound implications in society.

1. Morris, J. A., Gobrogge, K. L., Jordan, C. L. & Breedlove, S. M. 2004. Brain aromatase: dyed-in-the-wool homosexuality, Endocrinology volume of a sexually dimorphic nucleus in the ovine medial preop- 5. Roselli CE, Larkin K, Resko JA, Stellflug JN, Stormshak F. 2004. The tic area/anterior hypothalamus varies with sexual partner prefer-

145:475–477. 6. James, W. H. 2005. Biological and psychological determinants of male 2. Breedlove, SM. 1992. Sexually dimorphism in the vertebrate nervous ence. Endocrinology 145:478-483 - and female human sexual orientation, Journal of Biosocial Science system, The Journal of Neuroscience 12: 4133-4142. 3. Levay, S. 1991. A Difference in Hypothalamic Structure between Het - erosexual and Homosexual Men, Science 253:1034-1037. 37:555–567. tive pheromones in homosexual men, Proc. Natl. Acad. Sci. USA 4. Allen LS, Gorski RA. 1992. Sexual orientation and size of the anterior 7. Savic, H. Berglund and P. Lindstrom. 2005. Brain response to puta commissure in the human brain, Proc Natl Acad Sci USA 89:7199– 7202. 102:7356–7361.

36 | THE NERVE REVIEWS

Huemer’s Theory of Perception: Analysis and Objections By Ethan Rubin

SPRING 2010 | 37 REVIEWS

n his book Skepticism and the Veil of Perception, prehension, therefore, must represent an object as Michael Huemer lays out an account of perception actually existing, as opposed to representing it as a Ithat supports a version of direct realism. He states possibility or command. two main theses: that perception is direct awareness Huemer also requires that the apprehension of an of external reality, and that it leads to non-inferential object at least roughly correspond to the nature of the knowledge of that reality. The second claim requires object. He calls this “satisfying the content of the representation”. By accepting the possibility of some de- of Chapter IV in particular. In this paper, I intend to viation, he allows for certain common illusions, such mapthat theout first Huemer’s be adequately argument defended, and assess which its isstrengths the goal as the appearance of a bent stick in the water. The and weaknesses; each section explaining a key point correspondence between the awareness of the stick in his theory. After it has been explained, I will bring and the stick itself is close enough that the viewer can up objections to certain aspects of the theory as they correctly identify it and describe it with reasonable appear in the text, and consider the most promising accuracy. If the stick were perceived as a different ob- defenses against them. ject, such as a green kitten, there would not be enough correspondence between the apprehension and its object to consider the relationship awareness. Awareness requires an apprehension and a non- Huemer also stipulates that the correspondence accidental correspondence between apprehen- between apprehension and its object cannot be acci- sion and object dental, stating that “the apprehension must have been formed in such a way, and under such conditions, as to - make its correspondence with reality probable”. For ing direct awareness. He uses the term “aware- instance, if one were to hallucinate a particular scene Hness”uemer’s in firsta technical task is tosense, clarify as his the claim relation regard be- while that scene coincidentally occurs elsewhere, he tween one who is aware and the object of which he or she cannot be said to be aware of that event. Note is aware. This establishes that one must be aware of that this scenario is unsatisfactory even if the correspondence is perfect. This sort of example highlights cannot be actual awareness. This excludes cases of two problems with accidental correspondence: the hallucinationsomething – iffrom there awareness. is no object present, then there apprehension is not of the true object, and the observ- Huemer claims that awareness must include mental state of assertive representation. Not all men- er does not have sufficient reasons to believe that it is. tala state states, of heapprehension. claims, have He representational defines this term content as a Direct awareness is unmediated (the example he uses is the sensation of tickling). Thoughts, desires, and perceptions, however, do have uemer’s theory hinges on the difference be- representation content, but represent their objects tween direct and indirect awareness. In indi- in different ways. Apprehensions are characterized Hrect awareness, he states, one’s awareness is by actualized representation, meaning that they rep- of x, but only by means of his awareness of something resent their objects as being the case. He calls this else. When driving a car, for instance, one is aware of characteristic the apprehension’s assertiveness. Ap- the amount of gas left in the tank. This awareness is

38 | THE NERVE REVIEWS indirect because it is based on the driver’s awareness Huemer recognizes another sort of perceptual er- of the indicator needle on the dashboard, which he or ror that must be explained. It is possible to perceive - an object in a manner that corresponds to the actual rect awareness, by contrast, is unmediated by such a nature of the object and still mistake it for something secondaryshe expects awareness. to reflect the In amountorder to of be gas indirectly in the tank. aware Di other than it is. For instance, one might see a coil of rope and mistake it for a snake. To resolve this, Hue- of something else. Indirect awareness is a causal and mer introduces the notion of primary versus second- of something, one must first have a direct awareness ary perception. In secondary perception, the object is awareness by a logical connection between it and the perceived by virtue of another perception: perceiving logical relationship – one is led to reach the second the coil of rope or the snake involves the same pri- awareness to the indirect. This requires that direct mary perception, as the same image is seen in both awarenessfirst. As such, refer it transmitsreliably to theexternal authority reality. of the direct cases. The difference lies in the secondary perception, Three Elements of Perception: a Perceptual Expe- in which that same image is taken to be of different rience, an Object of Perception and a Causal Rela- objects. The coil of rope is therefore seen as a snake. tionship This sort of error is only admissible to a point, much like the case of illusion in primary perceptions. A mis- Having laid the foundation for his theory, Huemer taken secondary perception only arises from a primary perception that has a reasonable resemblance major components of perception, each structured to the secondary perception; a coil of rope does have similarlyturns to an to analysisawareness. of perception. First, there He is identifiesa purely interthree- some visual resemblance to a snake. If someone were nal mental state, which he calls the perceptual experi- to mistake the coil of rope for a bear, there is a more ence. Second, there is an object of perception, which serious error at work. Finally, there must be a connection between the of the experience. Third, there is a causal relationship internal experience and the external object. Huemer betweenis external the and two. at least The absenceroughly satisfies of any of the these content ele- proposes that the object must have caused the perceptual experience if the experience is of the object, perception. thereby excluding coincidental correspondence. This mentsThe disqualifies perceptual the experience event in differsquestion from from percep being- causal relationship also must be direct; Huemer ex- tion as a whole in that it does not necessarily entail an cludes what he calls deviant causal chains. If you were external object. Because the experience is internal, it to have an accurate perceptual experience, but only is not necessary that it be accompanied by an object. because someone with the appropriate knowledge This is how Huemer accounts for hallucination. A per- implanted the perception in your mind, it is true in a ceptual experience does occur in hallucination, but sense that the presence of the object caused the per- there is no external object present. The perceptual ceptual experience. The causal chain, however, has an experience still occurs because on an internal level, intermediary link and is not perception. hallucinations are indistinguishable from genuine perception. This implies that they share a common mental state and that internal experiences should be Three Elements of Perceptual Experience: Senso- recognized as separate from external objects of per- ry Qualia, Representational Content and Forceful- ception. ness As in awareness, the object of perception must meet a standard of resemblance to the content of the - perceptual experience. This standard allows for some ponent of perception, perceptual experience, discrepancies, but only to a point: when the experi- Hintouemer three proceeds features. to Hesubdivide claims thatthe first a percep com- ence represents the object as being fundamentally tual experience always has sensory qualia, which are different than it is in reality, it can no longer be called perception. The content satisfaction criterion is pres- also must have representational content, and that ent in other representations as well. A painting, for defined as “what it is like” to have the experience. It example, can diverge from its subject’s appearance to seeming present and real. These components are no- a certain extent, but at some point it can no longer be tablycontent similar must to have those forcefulness of awareness. – the characteristic of considered a painting of that subject. According to Huemer, all experiences are accom-

SPRING 2010 | 39 REVIEWS panied by qualia; sensory qualia are those that corre- is propositional. A perceptual experience expresses spond to perceptual experiences, as opposed to emo- the proposition “it is true that object X is before me,” tional or imaginative experiences. Qualia exist over which is a precondition for reaching conclusions re- and above representational contents, and are ineffa- garding that object. This proposition leans on another, ble in that they cannot be explained to someone who namely that “it is true that my perceptual experience has never had a comparable experience. Someone portrays the external world with satisfactory accu- who was born deaf, for example, cannot understand racy.” what it is like to hear. This ineffability is not mystical Huemer is careful to point out, however, that propositional content does not necessarily imply con- “...we have knowledge via ceptual content. Conceptual content, unlike representa- perception, which is not tional content, is not an intrinsic property of an expe- the same as saying that all rience. A perception is the same whether the perceiver perceptual experiences has concepts for all the objects, for only some of them, or for none at all. In fact, we constitute knowledge.” cannot have enough con- or metaphysical, nor does it apply to common experi- cepts to address the nuance and variability of experi- - impaired hearing can describe sounds to each other differentiate between the many shades of red that we effectively.ences and normal perceivers – two people with un perceiveence. For as example, being distinct. the concept The use“red” of isdemonstratives insufficient to On Huemer’s theory, perceptual experiences also can address them because it has a pointing function, involve representational content. He explains that saying “I see it is that, or thus and so.” This pointing “things appearing to be a certain way is not some fur- does have propositional content, i.e. “it is true that I ther consequence of your experience; things appear am perceiving that object, which is that way,” but the a certain way by virtue of your having the perceptual perceiver does not have concepts for “that object” or experience itself”. This assertion challenges theories “that way.” of perception that ascribe appearances to an act of By rejecting the necessity of conceptual content, interpretation that takes place after the perceptual Huemer is not rejecting the possibility of conceptual experience. For Huemer, representational content is content. He agrees that conceptual content exists and an essential part of the experience. Consider the bent is capable of affecting perceptual experience. His point stick illusion: the stick is not interpreted as being bent. is that perceptual experiences can occur without con- On the contrary, the experience represents it as being ceptual content, implying that conceptual content is bent and the viewer interprets this appearance as an not an intrinsic or essential component of perception. illusion. If interpretation determined the object’s ap- He refers here to Wittgenstein’s ambiguous “duck- pearance, the illusion would only occur if the perceiv- rabbit”. He admits that if one has the concepts of duck er believes the stick is bent. But no normal perceiver and of rabbit, the picture has the conceptual content would argue that the stick is bent, despite the fact that of both duck and rabbit. If one only has the concept it appears to be so. The temptation to say the stick is of duck, however, the picture will not have the con- bent is based on how the stick looks. Hence, this ap- ceptual content of rabbit. If the viewer does not have pearance must be an integral part of perceptual expe- either concept, the picture has no conceptual content, rience and independent from interpretation. but the viewer still has a perceptual experience. Per- Representational content is crucial to learn- ceptual experience can be altered by the inclusion of ing because it represents the object of perception as concepts, but it does not have to be conceptualized something that actually exists. This evokes Huemer’s and does not depend on conceptual content. - The propositional, assertive nature of perceptual ence represents its content as actualized, or as being experience leads Huemer to his discussion of forceful- thedescription case. It followsof assertiveness is that representational – a perceptual content experi ness. He begins by asking how imagining and perceiv-

40 | THE NERVE REVIEWS

imagines a duck or sees one, the content is a duck in Perceptual experience cannot be caused by mental bothing differ. cases. The One two possible can have difference the same is content that a percep– if one- orhave brain the states perceptual because experience they do not of bear synapses resemblance firing. tual experience has more detail and precision than an to its contents. The visual (or auditory, olfactory, etc.) imaginative one, but this difference can be overcome by focus and training, or by a photographic memory. experiencing. Detail, therefore, cannot be the fundamental distin- experienceBased onis first these and arguments, foremost of Huemer the object concludes one is guishing factor between the two. No one confuses that perceptual experiences are directly caused by imagining with perceiving, because the latter rep- physical objects that have the attributes that the expe- resents its content as being actualized, whereas the rience represents them as having. According to Hue- former does not. Huemer calls this difference force- mer, it follows that “In the primary sense of ‘aware’… fulness. This should not be confused with Hume’s we are directly aware of the fact that there are objects concept of vivacity, which is more like the difference with those colors and shapes”. That is, because our in detail that Huemer considers and discards. Hume’s perceptual experiences are direct, the knowledge we vivacity is merely a question of faintness or vividness, gain from them is also direct. He attributes the oppos- while Huemer’s forcefulness contains an element of ing argument, which holds that knowledge of exter- being present. nal objects is indirect because it is based on perceptual experiences, to a confusion between the object of awareness and the vehicle of awareness. This is clari- Why This Is Direct Realism chop wood. He is not, however, chopping the axe, but ne could support the majority of Huemer’s fied by the following analogy: one must use an axe to claims and still be a proponent of indirect real- the object of the chopping and the axe is the vehicle, Oism by arguing that perception makes us aware oris chopping means, of the the wood chopping. by using Applying the axe this – the to wood percep is- of mental states that become knowledge by a second- tion, the perceptual experience is the metaphorical ary, non-perceptual process. Huemer, however, insists axe that “chops” external objects. We cannot perceive that we are directly aware of more than just mental an object without having a perceptual experience that represents it to us, but that experience is only a tool. The awareness is directly of the objects by means of awarenessstates and thatconsists his theoryof an apprehension, of perception orsatisfies assertive his perception, just as wood is chopped by the man using mentaldefinition representation. of direct awareness. His description As he explained of perceptual earlier, the axe. - To put it another way, we do not perceive our sion: perceptual experiences have forcefulness (mak- perceptual experiences. If awareness were based on ingexperiences them assertive), accords qualia with the (making definition them of mental) apprehen and perceptual experience, we would have awareness of representational content. external objects by perceiving experiences, but this is not the case. We perceive external objects by hav- awareness by stipulating a causal connection between ing perceptual experiences, not by perceiving percep- the perceptualHuemer’s theoryexperience fits and the the definition object of of percep direct- tual experiences. The same goes for awareness: our tion. Saying that awareness must be of something is awareness does not arise from being aware of an ap- equivalent to saying that it must be caused by its ob- prehension, but simply by having an apprehension. ject. Therefore, asking what a perception is awareness Perceiving perceptual experience or being aware of of is also asking what causes the perceptual experi- apprehensions requires a second order act in which ence. Huemer maintains that only physical facts can one turns his attention to processes that occur wheth- satisfy the contents of perception and therefore must - be their source. It makes no sense to say that men- quiring introspection and deliberate effort, not a con- tal states cause perceptual experiences because the stitutiveer or not elementhe reflects of perception.on them. This is an extra step, re contents of perceptual experiences are not present in mental states; mental states do not have the properties, such as shape, color and texture, which form the contents of perceptual experiences. He rejects brain states for the same reason; normal perceivers do not

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Objection 1: How to Recognize Knowledge as well. The mental state involved in hallucination is - uemer fails to account for a practical problem tion is subject to review over and above the experi- in his theory. He admits that one must be ca- encethe same itself, as normal that involved perception in perception must be subject– if hallucina to the Hpable of handling certain mishaps that tend same process of judgment. The inclusion of judgment to befall perception from time to time, but does not in this context begins to stray dangerously close to in- make any conclusive statements about how this is direct realism, which Huemer wants to avoid. to be done. Hallucination and correctness by coinci- Similar arguments can be made regarding Hue- dence are problems that he needs to address more mer’s demand for a direct causal connection between thoroughly. apprehension or perceptual experience and the object represented. The distinction he makes between relationship and therefore must be of something. This an acceptable connection and an accidental or deviant conditionHis definition excludes of hallucination awareness stipulates from awareness that it isbe a- one is once again external rather than internal. One cause one cannot have a relationship with something example he uses of accidental correctness is that of that does not exist. There is an obvious external dif- calling a friend. *After the phone rings multiple times, ference between awareness and hallucination that one may conclude that the friend is not home. How- makes them fundamentally distinct, but this does not ever, one could unknowingly have dialed the wrong suit Huemer’s project. Since he is describing aware- number and not actually tested whether the friend ness phenomenologically, from the inside, he must was home or not. The conclusion may still be correct account for how the perceiver himself understands by coincidence and, by asking the friend whether he his mental states. It makes sense to say that one can- was home at the time, he can come to know that his not have a relation with a nonexistent object, but if he conclusion was correct, and yet this does not qualify feels like he is in a state of awareness and cannot dif- as awareness. The reasons why it cannot be aware- ferentiate between his hallucination and a real object, ness are clear to the third party reading the example, it is unclear how the distinction is salient. but the person in the example has no access to those Huemer does admit that there is some irregular- reasons. There is no way for him to verify the relation- ity in perception. His position is that we have knowl- ship between his experience and his apparent knowledge via perception, which is not the same as saying edge. that all perceptual experiences constitute knowledge. A deviant causal chain presents the same prob- Even so, the fact that we cannot tell the difference be- lem. In Huemer’s example, someone has a perceptual tween awareness and hallucination is problematic be- experience of a cup. A scientist has implanted an exact cause it impedes knowing which experiences lead to copy of the perceptual experience caused by the cup in knowledge. Huemer could argue that one is inclined the would-be observer’s brain, with the result that the to judge that an experience is hallucinatory when it experience does correspond with the object it repre- does not accord with his expectations of reality, cor- sents. The cup caused a perceptual experience in that recting for errors in perception after the fact, but this - raises two problems. First, it is only a plausible solu- ence, but the subject cannot be said to have perceived tion if the hallucination is recognizably outlandish theit caused cup because the scientist the causal to implant connection that specific is too experifar re- and depicts an “object” whose existence is improb- moved. The reader has no trouble understanding why able enough to make the hallucinator disregard his this is not perception, but the deceived character in the example cannot reach the same understanding. In state is the same in both perception and hallucination, light of these examples, causal connection seems like theperceptual only difference experience being – rememberthe presence that or theabsence mental of a dubious criterion. If one cannot distinguish between the object. It is not implausible that one could have the causal chain of a genuine apprehension and that a mundane hallucination of an unsurprising object, of an “apprehension” based on coincidence or decep- or even an existing object that he had accurately per- tion, then one cannot use causal connection as a stan- ceived before. There would be no cause for suspicion dard for accepting or rejecting his own experiences. and he would accept the hallucination as awareness. It seems that Huemer owes us a better solution to the Second, inserting an act of judgment in the case of hal- internal problems of recognizing knowledge. lucination would leave Huemer open to the objection that such an act of judgment takes place in other cases

42 | THE NERVE REVIEWS

Objection 2: Where to Draw the Lines central to his theory, how can Huemer justify glossing over what a deviant causal chain is? He is obligated

should make the reader wary. By making pro- that the deviant chain is “convoluted and abnormal” Hvisionsuemer is for satisfied rough correspondencewith some ambiguities between that a replacesto specify “deviant” what is deviant with synonyms and what and is direct does –nothing saying perceptual experience and its object but disqualify- to pin down what it means. ing those that do not correspond enough, he commits This problem could also be addressed by appeal- himself to a spectrum that covers the entire range being to case-by-case decisions. For the most part, what tween exact resemblance and complete disjunction. At Huemer means by a deviant causal chain is under- some point in this spectrum, experience ceases to be perception and becomes hallucination. Most readers objection incites a sort of Wittgensteinian reply, as in would ask Huemer to indicate where this point lies. It “youstood know despite what the I absencemean, this of ais precise perfectly definition. clear to Theyou is easy to make an absurd example that highlights the when the case arises.” This solution works better here than it would for the previous objection. The resem- stick in water that appears bent and seeing a stick in blance condition must allow for a range of illusions thedifference water that between appears the to extremes be a green – compare kitten. The seeing closer a and imprecisions, whereas the causal connection is a we come to the middle of the spectrum, however, the stricter requirement. The presence of virtually any in- termediate cause between the experience and its ob- stick in water that appears bent and seeing a stick in watermore difficultthat appears it is toslightly distinguish more bent – compare than the seeing optical a makes it much easier to consider a case and decide phenomenon accounts for. How much more bent can thatject isthe sufficient relationship to disqualify is not direct. the causal chain, which the stick appear before one is no longer perceiving the stick? Huemer might respond that the spectrum is am- Objection 3: Unnecessary Components of Percep- biguous by nature and that distinctions must address tual Experience the case at hand and its context. This may be true, but there must be some general standard in place by uemer’s analysis of perceptual experience which to make the distinctions in each case. Even if it does a good job of analyzing the various fea- highlights a vague section of the spectrum as the area Htures that are present, but his insistence that in which experience begins to lose legitimacy, there they are separate entities may be misled. He claims should be some kind of criterion. Saying that accept- that sensory qualia, representational content and able cases “roughly satisfy” the representation and forcefulness are all different components of a per- unacceptable ones “diverge radically” tells us very ceptual experience. Upon consideration, however, it seems that they are inextricably linked and cannot the intrinsic fuzziness of the correspondence condi- be disjoined. Rather than split up perceptual experi- little about what qualifies and what does not. Despite ence into three elements, it would be more accurate to the spectrum and identifying the boundaries of per- speak of it as a single entity with features that account ception.tion, Huemer could stand to do a better job of defining for its particular nature. Forcefulness, for instance, He falls prey to the same pitfall in describing the can be considered an element of representational deviant causal chain. He states that the causal con- content, as it is essential to and inseparable from per- nection between perceptual experience and its object ceptual experience. Without forcefulness, the experi- must be close enough that the object directly causes ence is no longer a perceptual one: it is an imagina- the experience, but “directly” does not specify what tion or a memory. This is because forcefulness is a feature of how the object of perception is represented sentence, “I shan’t enter into the question of how ex- to the perceiver, namely as being present. But Huemer qualifies. I had to suppress laughter upon reading the- had posited assertiveness long before he introduced forcefulness. Assertiveness is an essential aspect of componentactly one might of perception, define ‘deviant in whose causal absence chain’….” percep Hue- representational content and lends immediacy to ex- tionmer identifiescannot occur direct at causal all. A connectiondeviant causal as an chain essential pre- perience; seeing as assertiveness does the same work cludes direct causal connection, meaning that the ex- as forcefulness, there is no reason to tack forcefulness perience in question is not perception. Seeing as this is onto the end of the theory.

SPRING 2010 | 43 REVIEWS

Sensory qualia are the most dubious of the com- qualia because we no longer have the sense that the color we experience is something strange and illegiti- with the disclaimer that their ineffability is not mys- mate that must be explained away. teriousponents and Huemer must notidentifies. be mistaken He begins for “a his resort discussion to ob- Upon asking whether there is any reason for qua- scurantism and mysticism”. This is not, however, the lia to exist or if they serve any purpose, Huemer exhib- strongest objection against qualia. Even having ac- its more unintentional humor by responding “I do not cepted their ineffability as a characteristic of experi- know the answer to this”, yet still insisting that they must exist. He offers two potential answers, neither Because perceptual experiences are internal, as Hue- of which is very convincing. First, he claims that the merence, states, the existence qualia could of qualia just asstill easily seems be superfluous. an intrinsic information obtained through perceptual experiences characteristic of perceptual experiences without be- could not be represented without qualia. The very fact ing a separate class of mental entities. that he uses the word “represented” betrays his ar- Huemer’s point is that the representational con- gument. He asks how we could perceive red without qualia and swiftly concludes that it is impossible, but because there is some difference between the object it is perfectly plausible that we perceive red by way andtent theand experience,the object thatsome satisfies “what it areis like” the tosame, experi but- of representational content, which represents the ob- ence the object, qualia must exist. The function he ject as being red, and therefore have no reason to posit attributes to qualia, however, can easily be included qualia. Second, he asserts that qualia serve a conative in the notion of representational content. Representa- function, expressing the pain or pleasure of an experi- tional content represents an object as being a certain ence. There is no obvious reason why representation- way to the perceiver. This is an aspect of his internal al content should not perform this function as well, mental state and as such has the same ineffability as representing a warm experience as pleasant or a red qualia; it represents the object as being “this way.” As experience as striking. He argues that qualia “simulta- it represents the object to the perceiver, the perceiver neously give us information about the world and give experiences the representational content. How is this us emotional reactions or desires. Those two func- different from qualia, “what it is like” to perceive an tions are integrated into the same experience, rather object? The distinction implies that representational than being functions of two separate states or events”. content is a component of perceptual experience that It is the purpose of representational content to give is not experienced in any particular way, which is ab- us information about the world; if the two functions - are integrated, it makes more sense to simply assign perience is what it is like to have its object represented them to representational content and discard qualia tosurd him. – “what How elseit is like”could for representational one to have a perceptual content pos ex- as redundant. sibly function? There is no way that content could be represented to the perceiver except by his experiencing it, nor is there anything outside of his experience Objection 4: Sense Data and Representationalism that he is aware of when he perceives something. The nature of representational content thus neutralizes hen he explains how his theory is a form the need for qualia to explain experience. of direct realism, Huemer makes a danger- Huemer’s explanation of the difference between Wous statement: “We might also be said to red and red* (the quale) is based on a confusion. He be aware (directly and primarily) of the colors and states that “…red* is a property of experience, where- shapes of the (facing surfaces of) physical objects as red … is a property of physical objects”. It is correct around us, since that could also be described as what to separate these as properties of two different types, but the way in which he speaks of them is mislead- be objects of those colors and shapes”. Although he ing. Red as a property of physical objects is not a color, describessatisfies the this content view as of if ita werestate onlyrepresenting a trivial variation there to of his previous claim, it opens the door to a host of op- of light. Color, on the other hand, is an experienced posing theories that he wants to avoid. In particular, property.but rather It the would tendency be more to reflect logical certain for red* wavelengths to refer to it allows for the possibility of a sense datum theory, which in turn implies representationalism. indicate the color that is experienced. From this per- He ends this section by saying “We must therefore spective,the physical it isproperty unnecessary of reflecting to posit light the and existence for red to of conclude that direct realism is true”. This conclusion

44 | THE NERVE REVIEWS follows from his original claim that we are directly objection three is the most radical, but Huemer could aware of the existence of objects, but it does not fol- address it easily. Neither the existence of qualia nor low from the sense datum theory he implies in his the independence of forcefulness is so central to his second claim. According to the latter theory, we are argument that his theory would be irreparably dam- directly aware of sense data, not objects. In order to aged by removing them, and all the phenomena they have awareness of objects, one must undergo a non- explain could be preserved in the updated account. perceptual process of judgment, forming a theory that Objection four, if correct, deals a heavy blow to infers the existence of objects from the sense data. Huemer’s original theses. Sense datum theory would Therefore, sense data do “satisfy the content of a state representing there to be objects”, but they do not do so awareness of sense data. The second thesis would directly because the state is based on the sense data. thenrewrite have his to first be thesisrevised to to say say that that perception we have indirect,is direct By admitting the plausibility of this view, Huemer is in inferential knowledge of the external world as a result danger of committing himself to indirect realism. of the sense data gathered by perception. Although The axe and wood analogy Huemer provides is Huemer only becomes explicitly vulnerable to sense clever and useful, but it does not grant him immu- datum theory at the end of the chapter, the rest of his nity from sense datum theory either. He intends the account would not need extensive alteration to ac- physical objects to be the “wood” and perceptual ex- commodate sense data. perience the “axe”, but the analogy works equally well If we replace his references to physical objects if we replace “physical objects” with “sense data.” with references to sense data, we arrive at the fol- The argument would go as follows: We use percep- lowing theory: perception consists of an internal tual experience as a tool that enables us to perceive mental state, an object that produces sense data in sense data directly. As a consequence, we are aware of the perceiver, and a causal relationship between the physical objects only indirectly. Huemer could object mental state and the sense data. The internal mental that sense data are also vehicles of perception, but his state consists of the experiences of perceiving sense analogy betrays him again. If perceptual experience data (whether or not these are called qualia), whose is the vehicle of perception, sense data must be the contents represent patches of color as immediately objects of perception. Once the sense data have been present (whether or not this is called forcefulness). perceived, the analogy repeats: sense data become the Therefore, perception is direct awareness of sense vehicle by which we become aware of physical objects. data, which leads to indirect knowledge of the exter- At this point, however, awareness has already become nal world by a process of judgment. indirect. In order to use them as means, one must This is by far the greatest threat to Huemer’s proj- - means, they are not vehicles of perception but rather first obtain the sense data as objects. When used as forect him– it challengesto argue against. his central One thesesdefense without Huemer contra could occurred. Physical facts, in turn, become the objects mountdicting isthe that majority we are notof his aware theory, of any making process it difficultof judg- of judgment judgment – and all the are necessary thus distanced perception from has the already direct ment between perception and awareness. If sense relationship with perception that Huemer wants to data require judgment to be seen as objects, why do maintain.

Conclusion theory,we see however,objects in lies the with world Huemer without himself. having to reflect on them first? The task of filling the breaches in this or the most part, Huemer’s theory is plausible REFERENCE and supports his theses. Objections one through Fthree do have some validity but are not strong Huemer, Michael. Skepticism and the Veil of Percep- enough to falsify his system in general. Some can be defended against, while others can be reconciled to his 2001. tion.Oxford: Rowman and Littlefield Publishers, Inc., account with only minor revisions. Of the first three,

SPRING 2010 | 45 OPINION

Homemade Terror

“Fear is a tyrant and a despot, more terrible than the rack, more potent than the snake.” ~ Edgar Wallace

“People react to fear, not love; they don’t teach that in Sunday school, but it’s true.” ~ Richard M. Nixon

and no one knows this fact better Who can forget than media corporations and poli- frenzy. the emotions that burst from the ticians. The Patriot Act, signed by AmericanA radio anxiety address into given a post by Presi 9/11- American psyche in the period President George W. Bush October dent George W. Bush on March 22, following September 11, 2001? 26, 2001, legalized roving wire Bewilderment, despair, outrage, taps, greater access to individu- “The people of the United States and a profound sorrow gripped als’ private records, and “sneak & and2003 our proclaimed friends and the allies following, will not our collective consciousness as we peak” warrants. Most were willing live at the mercy of an outlaw re- watched the symbols of our coun- to accept these policies, as long gime that threatens the peace with try crumble in dust clouds that only as they guaranteed their families weapons of mass murder. Now that temporarily obscured the grim re- could sleep safe from the ever pres- ality that waited to be unearthed. ent specter of terrorism. limit its duration is to apply deci- As America reeled from this trau- siveconflict force. has This come, will the not only be away cam to- matic blow, one reaction lingered force headed by the United States like the scent of rotted meat. We and OnEngland March began 20, 2003 an aptly a Coalition named for the security of our nation and became a nation united in fear. “Shock and Awe” bombing of Bagh- thepaign peace of half-measures. of the world, and It is we a fight will The media splashed images, dad. These explosions heralded an accept no outcome but victory.” of snarling faces pledging our de- invasion and eventual occupation America was ready to initiate struction, upon our mental can- that will continue into the foresee- war to secure the peace. But as al- vases in what seemed like a non- able future. In the months of debate legations of deliberate twisting of stop delude. In the weeks following information mounted, support for such as Al-Qaeda and weapons of the war plummeted. According to of terror launched not from over- massleading destruction to the first were strike, showered phrases a CBS News/New York Times poll seas9/11, but we from suffered within from our a campaignown bor- upon the American public. These the percentage of Americans who ders. How much of this fear satura- buzzwords in conjunction with the thought the “war with Iraq was tion was used to further economic color-coded terror threat levels, worth the loss of American life and and political agendas? Fear sells, enacted March 11, 2002, pushed other costs of attacking” dropped

46 | THE NERVE OPINION

odor of a predator) are more likely cognitively changing the way we in- to escape threatening situations, terpret a situation, we can actively dissipatedfrom 46% on so August, did America’s 2003 to disre 24%- alter the way we feel about it. gardon September, for the costs 2009. of war. When the fear who don’t. Imagine you’re about to de- What is it about fear that al- and Imagestherefore that more inspire ‘fit’ than fear, those such liver a speech in front of an audi- ence (public speaking may be one our thoughts and actions? Discov- Towers and Osama Bin Laden with of our biggest fears). You can feel erieslows it in to Cognitive so drastically Neuroscience influence threatsas the flaming streaming collapse from ofhis the lips, Twin can your anxiety rise and heart race as may hold some of the answers. you envision possibilities of fail- One of the most provocative were not consciously paying at- ure and embarrassment. One way tentioninfluence to ourthem. behavior Think backeven towhen the many combat this negative affect are processed automatically and is through reappraisal. They tell subconsciously.findings is that emotional Research stimuliby Dr. or to the period leading up to the themselves that they’re not ner- Kevin Ochsner and colleagues invasionmonths and of Iraq. years The following media 9/11 was vous - they’re excited and that this has found that emotional stimuli pervaded with images and stories excitement will lead to a better per- masked from conscious percep- of terror. How much did this bom- formance. They remind themselves tion still resulted in an increased bardment dictate our willingness that they have felt this way before autonomic response (i.e. increased to establish “the security of our na- and have still delivered. Reapprais- heart and respiration rate) and tion” no matter the cost? al has been found to be more effec- amygdala activation. The amygdala Luckily, there is research into tive than both suppression and ac- is an almond-shaped collection how people cope with fear and ceptance in regulating the negative of nuclei found within the medial anxiety. The consensus emerging aspects of fear. temporal lobe. The amygdala is from the literature is that the best If more people had stepped thought to evolutionarily predate strategy for regulating anxiety is back and revaluated the informa- brain regions such as the neocortex reappraisal. Emotional states stem tion that was used to justify the in- (whose size and complexity sepa- from our evaluation of an emo- vasion of Iraq, war may have been rates human cognition from other tional situation’s context and ele- avoided. Maybe not. Either way, I animals’). From an evolutionary ments as they directly relate to us. am wary of those who would ma- point of view, organisms that have Shark attacks are more fear induc- nipulate my innate fear response to - ing than car accidents because of - tion to fearful stimuli (such as the the way we gauge the scenarios. By iors. Are you? an immediate and reflexive reac influence my thoughts and behav — Darrien Garay What is needed, rather than running away “ or controlling or suppressing or any other resistance, is understanding fear; that means, watch it, learn about it, come directly into contact with it. We are to learn about fear, not how to escape from it. ~ Jiddu Krishnamurti ”

SPRING 2010 | 47 Don’t toss out that term paper! Submit it to The Nerve!

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The Undergraduate Program in Neuroscience is an interdisciplinary major leading to a Bachelor of Arts in Neuroscience. As a field that has grown considerably through integration across disciplines over the last few decades, a current understanding requires knowledge that spans traditional approaches. The new program combines breadth of exposure to the field with the opportunity for depth of experience in one of three central domains of neuroscience: Cellular and Systems, Cognition and Behavior, and Computational Neuroscience. Students will have access to the extensive resources and expertise of faculty across multiple departments and colleges throughout the university. A wide array of courses are offered through the departments of Biology, Cognitive & Neural Systems, Computer Science, Mathematics & Statistics, and Psychology. Opportunities for independent laboratory research are available through multiple departments in the Colleges of Arts and Sciences and Engineering, and at Boston University School of Medicine, including Anatomy and Neurobiology, Biochemistry, Neurology, Pathology, Pharmacology and Experimental Therapeutics, Physiology and Biophysics, and Psychiatry.

The program is directed by Professor Howard Eichenbaum, University Professor, Director of the Boston University Center for Neuroscience, Director of the Center for Memory and Brain, and a Professor in the Department of Psychology. The academic director for the program is Dr. Paul Lipton, Director of Undergraduate Academic and Research Affairs, and Research Assistant Professor in the Department of Psychology. The program administrator is Lindsey Clarkson. Our offices are located on the first floor of 2 Cummington Street, Rooms 109 and 114 (Ph: 617-358-5150 or 617-358-3298). An advisory curriculum committee charged with overseeing curricular development and facilitation of undergraduate research opportunities is composed of Professors Vincent Dionne (Biology), Nancy Koppel (Mathematics and Statistics), Ennio Mingolla (Cognitive and Neural Systems), David Somers (Psychology), and Paul Lipton (Psychology).

All courses available for credit toward a B.A. in Neuroscience are taught by affiliated faculty housed within multiple departments in the College of Arts and Sciences, including Biology, Cognitive and Neural Systems, Computer Science, Mathematics and Statistics, and Psychology, and Health Sciences in Sargent College. Together they offer over 50 upper level neuroscience electives, including laboratory courses and seminars. Undergraduate research opportunities are available in neuroscience laboratories throughout the university in over 30 departments across both the Charles River and Medical campuses.

Academic advising for neuroscience students is arranged and available through Dr. Paul Lipton. To set up an appointment, please email Lindsey Clarkson at [email protected].

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