DD303 TMA04 Stephen Lissenden Personal Identifier: W8607760

Using examples discuss what can be learned about normal cognitive function by using haemodynamic imaging techniques such as fMRI and PET on healthy volunteers.
Haemodynamic imaging uses noninvasive techniques to construct images of the brains metabolic activity and blood flow thus enabling inferences to be drawn in support of specific theoretical models of cognition. However, whilst this methodology has generated a large amount of interest, the extent to which such techniques can contribute to the understanding of cognitive function is debatable. This essay will present some haemodynamic imaging studies, the conclusions researchers have drawn and some alternative views of the evidence. Additionally a discussion of the underlying methodological and theoretical assumptions will examine the extent to which haemodynamic imaging may be useful in understanding cognitive function. Firstly, however it will be useful to briefly outline two types of haemodynamic imaging, the processes that enable them and the form of data these techniques provide. The brain requires oxygen and glucose to function and, as neural activity increases, blood flow to these areas of activation increases; delivering the additional oxygen and glucose that is required. FMRI measures changing oxygen levels, detecting what is known as the BOLD response, whilst PET, using radioactive tracers, detects both neurotransmitters, the chemicals that transmit signals between neurons across the synaptic cleft, and glucose. Using both fMRI and PET scans, inferences are drawn regarding the location of specific brain functions and, more importantly to cognitive psychologists, provide an insight, although indirect, into the implementation of cognitive process within the brain. PET scans can locate the position of neural activity to within 10mm whilst fMRI has a resolution of up to 1mm. However, whilst measuring location with relative accuracy, these techniques have a poor temporal resolution. In the case of fMRI it can take up to five seconds after neurons have fired for a BOLD response to be detected whilst PET scans can take anywhere from 10 seconds to 20 minutes to produce an image (Biomedical Imaging Research Lab, University of Southern California, 2009). These temporal limitations can be minimised by combining heamodynamic approaches with EEG, as demonstrated by Opitz et al. (1999), or MEG techniques that measure near instantaneously the electrical or magnetic properties of neurons as they fire. However, although EEG and MEG are useful for identifying when activity is occurring, on their own they could not identify where the activity is happening but although all neuroimaging methodologies have their limitations, many studies have employed haemodynamic techniques to investigate cognitive functions. Paulescu et al. (1993) used PET scans to investigate the changing activity patterns within the brain that occurred during phonological processing. Experimental participants performed matched tasks that differed in phonological content. Then, using a procees of subtraction, where data from one experimental condition is removed from a second experimental condition to leave the data of specific interest, Paulescu et al. concluded

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that activation associated with phonological storage occurred in the supramarginal gyrus whilst sub vocal rehearsal was associated with the Brocas area. Paulescu et al. presented these findings as support for the Baddelley et al (1984) model of the phonological loop. Koepp et al. (1998) also used PET scans, this time to study the release, and subsequent binding, of the neurotransmitter dopamine within the ventral striatum, an area of the brain believed to be involved in the appreciation of reward. Upon finding a correlation between performance level when playing a video game and dopamine activity Koepp et al. suggested a potential link between performance and reward. Klein and colleagues (1995), using fMRI, studied the brain activity of bilinguals whose native language was English and their second language was French. The participants in this study repeated or generated synonyms of stimulus words in both languages. For word repetition, an area associated with skilled motor action, in the left basal ganglia, was more active when repeating French words, possibly due to an increased cognitive requirement when pronouncing these words. When generating synonyms in both languages the prefrontal cortex showed activation. Klein, using subtraction methodology, concluded that this demonstrated that the prefrontal cortex is associated with semantic processing and the selection of responses. Also using fMRI, Devlin et al. (2000) investigated whether the brain showed different areas of activation when categorizing living things and inanimate objects. Devlin et al. found there to be no significant differences suggesting there is no functional difference between these forms of categorization. However, despite the many studies using PET scans and fMRI, Harley (2004) suggests that whilst exploring the architecture of the brain may be useful, he raises doubts that neuroimaging, such as that provided by haemodynamic techniques, can, on their own, be used to develop theories of cognitive function. A number of methodological issues and theoretical concerns lend support to Harleys (2004) opinion. Whilst haemodynamic techniques offer persuasive images of the brain, these images are constructed from data, gathered from indirect, low resolution, partial measurements of neural activity. This data has undergone multiple transformations, using concepts such as subtraction, standardised space and statistical thresholding, embedding and masking assumptions that may, or may not, be valid when drawing conclusions regarding cognitive functioning. The indirect nature fMRI and PET scans, where inferring neural activity from changes in oxygen levels, is likened by Hardcastle & Stewart (2002) to trying to ascertain, by merely measuring the overall oxygen consumption in a room of 1000 people, who is sleeping and who is exercising. Hardcastle & Stewart also suggests that these indirect observations are partial, missing aspects of cognition, such as changes in the synaptic membrane that impact the flow of information. Hardcastle and Stewart (2002) also argue that the comparatively low resolution of fMRI and PET scanners has led to a misplaced support for modularization. As the resolution of scanners increases, the number of processing sites being identified also increases, leading Hardcastle and Stewart to see cognition as a distributed process. Page (2006) seems to support this view, pointing out that any brain region supporting separate cognitive functions can contain neurons that have either no direct neural connections, or, have some overlapping connections, or indeed, could use all the same neurons, but in a multiple configurations.

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Hardcastle and Stewart underline this opinion, pointing to studies that identify the premotor cortex as being active during numerous cognitive functions. They also note that premotor cortex activity data, subtracted from some studies, further simplifies the complex nature of the region, thereby seemingly providing mistaken support for modularization. Page (2006) also suggests that spatial separation is often assumed to imply functional separation, an assumption that is undermined to an extent by Hardcastle & Stewarts (2002) assertion that all neurons are probably linked to each other by fewer than six connections. Hardcastle & Stewart also suggests that there is a tendency within neuroscience towards separation and modularisation, contending that not only is there no evidence to support this view and that current methodologies cannot provide the evidence but also that the tension between a modularised model of cognition and a multi-tasking, distributed processing perspective is holding progress back. The debate regarding single and dual-mechanism models of recollection illustrates this point. Henson (2005) assumes that imaging can be used to falsify one of these models, indeed Henson concludes the imaging data support dual-process models over single-process models. (Henson (2005) p. 201) However, Coltheart, (2002) (cited in Page, 2006) a supporter of the dual-route model within the context of reading, will not entertain imaging as support for his position because he does not believe that any imaging data could be used to falsify his position. The dual route model has both routes engaged during processing, but these routes could be in the same region and even if one route was not engaged, the suppression of this route may still generate a BOLD response. Additionally, if two spatially separate regions are active this does not necessarily mean two distinct processes are in play. Another concern is the use of subtraction. As outlined earlier subtraction is a process by which to compare neural activity between two or more experimental conditions. The baseline activity pattern in one condition is subtracted from the activity pattern associated with a second condition, leaving, it is assumed, a neural activity pattern specifically associated with the second experimental condition. There are some issues with this assumption however. Hardcastle & Stewart (2002, p.78) suggest it is not possible to differentiate between any concurrent, coincidental activity and the cognitive process under scrutiny and Page (2006) reinforces this view raising concerns that functional relevance may mistakenly be attributed to epiphenomenal activity. Logothetis et al. (2001) (as cited in Page, 2006), a pioneer and proponent of haemodynamic imaging, also highlights that statistical analyses and thresholding methods applied to the haemodynamic responses probably underestimate a great deal of actual neural activity again outlining the simplifying nature of imaging methodologies. A lack of standardisation between studies complicates and compounds this simplification. As Hardcastle & Stewart (2002) point out, researchers use different baseline conditions and different statistical methods for analyzing data. With different studies simplifying the data in different ways, not only are comparisons between studies difficult, but this also introduces an additional

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layer of subjective interpretation into images presented as objective representations of neural activity. Yet Poldrack (2006), has raised more concerns, calling into question logical inferences made when interpreting haemodynamic images, identifying in particular the method of reverse inference, by which the engagement of a particular cognitive process is inferred from the activation of a particular brain region. When a region of the brain activates during a task then, if other studies appear to show an association between the same region and a specific cognitive process, Poldrack suggests, an inference is sometimes incorrectly drawn that this task must engage that cognitive process associated with that region. This type of claim is false and potentially calls into question the claims made in the studies by Koepp et al (1998) and Klein and colleagues (1995) mentioned earlier. Page (2006) provides another specific example of flawed reasoning when he suggests that Henson (2005) may assume that memory strength correlates with the strength of related neural activity, an unsubstantiated assumption that not supported by evidence. Loosemore and Harley (2010) suggest that we are being flooded with accurate answers to questions about the brain location of mechanisms that we do not believe in and inaccurate answers to questions about the brain location of mechanisms that are currently not terribly interesting. Page (2006) goes further asserting that there are too many assumptions underlying imaging techniques to enable them to effectively differentiate between theories of cognitive function whilst Uttal (2001) dismisses imaging techniques as a modern day form of phrenology. Haemodynamic images of the brain activity are becoming more detailed and complex and whilst these images are visually compelling, their meaning is not clear. Even if fMRI or PET could identify where cognitive processes are performed, and it is currently uncertain that this could be the case, they cannot answer how, why, or even if, these patterns of neural activity cause cognition or give rise to observable behaviour. This question of how cognition occurs is important, where it occurs, is less important to cognitive psychology, after all most will agree that cognition takes place somewhere in the brain. In addition, the framing of cognitive models, with the necessarily limited and transformed data of haemodynamic imaging, may restrict the types of explanation possible and whilst such an approach may assist in falsifying erroneous hypothesis, the use of haemodynamic imaging could also lead to misleading oversimplifications. It is possible to construe this perspective as undermining a potentially useful tool in the endeavour of understanding cognitive function. The argument may also be framed as akin to saying that being able to see chromosomes and genes using a very powerful microscope does not improve the theory of genetic inheritance (Kaye, 2010 p.20). Whilst this defence of haemodynamic imaging may have some merit, it also remains true that whilst we can see synapses firing using a very powerful microscope, or in this case infer that they are firing using fMRI or PET scans, we are yet to see a meaningful thought.

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References Baddelley et al. (1984), cited in Kaye, H. (Ed.), Cognitive Psychology (2nd ed.) (p.165). Milton Keynes: The Open University, 2010
Biomedical Imaging Research Lab, University of Southern California (2009). Positron Emission Tomography Dynamic. Retrieved July 17, 2011, from http://neuroimage.usc.edu/ResearchPETDynamic.html

Coltheart, (2002), cited in Page (2006) What can't functional neuroimaging tell the cognitive psychologist?, Cortex, 42 (3), pp.428433 Devlin et al. (2000), cited in Kaye, H. (Ed.), Cognitive Psychology (2nd ed.) (p.19). Milton Keynes: The Open University, 2010 Hardcastle, VG. and Stewart, CM. (2002), What Do Brain Data Really Show?, Philosophy of Science, 69 pp. S72-S82 Harley, TA. (2004), Promises, Promises, Cognitive Neuropsychology, 21 (1), pp. 5156 Henson, R. (2005), What can functional neuroimaging tell the experimental psychologist?, The Quarterly Journal of Experimental Psychology Section A, 58:2, pp. 193-233 Kaye, H. (Ed.), Cognitive Psychology (2nd ed.). Milton Keynes: The Open University, 2010 Kaye, H. (Ed.), Cognitive Psychology Methods Companion (2nd ed.). Milton Keynes: The Open University, 2010 Klein (1995), cited in Kaye, H. (Ed.), Cognitive Psychology Methods Companion (2nd ed.) (p.39). Milton Keynes: The Open University, 2010 Koepp et al., (1998), cited in Kaye, H. (Ed.), Cognitive Psychology Methods Companion (2nd ed.) (p.44). Milton Keynes: The Open University, 2010 Logothetis et al., (2001), cited in Page (2006) What can't functional neuroimaging tell the cognitive psychologist?, Cortex, 42 (3), pp.428433 Loosemore, RPW & Harley, TA (2010), Brains and Minds: On the Usefulness of Localization Data to Cognitive Psychology. In M. Bunzl & S.J. Hanson (Eds.), Foundational Issues in Human Brain Mapping. Cambridge, MA: MIT Press. Opitz et al., (1999) cited in Kaye, H. (Ed.), Cognitive Psychology Methods Companion (2nd ed.) (p.35). Milton Keynes: The Open University, 2010 Page, M. (2006), What can't functional neuroimaging tell the cognitive psychologist?, Cortex, 42 (3), pp.428-433 Paulescu et al. (1993), cited in Kaye, H. (Ed.), Cognitive Psychology (2nd ed.)
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(p.167). Milton Keynes: The Open University, 2010 Poldrack, RA. (2006), Can cognitive processes be inferred from neuroimaging data?, TRENDS in Cognitive Sciences, Feb2006, Vol. 10 Issue 2, pp. 59-63 Thompson-Schill et al. (2002), Effects of frontal lobe damage on interference effects in working memory., Cognitive, Affective, & Behavioral Neuroscience, 2 (2), 109-120 Uttal, WR., The New Phrenology: The Limits of Localizing Cognitive Processes in the Brain. Cambridge, MA: MIT Press, 2001. Weber, MJ. and Thompson-Schill, SL. (2010), Functional Neuroimaging Can Support Causal Claims about Brain Function, Journal of Cognitive Neuroscience, Vol. 22 Issue 11, pp. 2415-2416, 2

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Stephen Lissenden 19th July 2011

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