Beyond dissociation: Exploring interactions between implicit priming and explicit recognition

Over the last 30 or more years evidence has accumulated in favour of the view that memory is not a unitary faculty; rather, it can be subdivided into a num- ber of functionally independent subsystems. Whilst dividing memory phenomena into these distinct subsystems has undoubtedly advanced our understanding of memory as a whole, the approach of studying subsystems in isolation fails to address potential interactions between them. Over the last few decades there has been a gradual increase in the number of studies attempting to move be- yond dissociation by characterising functional interactions between subsystems of memory. The main aim of this thesis was to contribute to this endeavour, by examining interactions between two specific subsystems that are positioned on opposite sides of the declarative and non-declarative divide in long-term mem- ory: priming and episodic recognition. Event-Related Potentials (ERPs) were employed to monitor neural markers of repetition priming and episodic memory during recognition tests with masked priming of test cues. In the standard pro- cedure, half of the studied and unstudied test trials began with a brief (48ms) masked repetition of the to-be-recognized word prior to the onset of test items; the remaining unprimed trials were preceded by the word “blank”. The pattern of priming effects across experiments was reasonably consistent, with differences between experiments directly related to the intended manipulations. In contrast to priming effects, the pattern of memory effects was variable across experiments, demonstrating that the engagement of explicit recognition signals is influenced by the outcome of implicit processing, and suggesting that interactions between priming and explicit retrieval processes do occur. Taken together, results from experiments reported in this thesis indicate (1) that under certain circumstances, priming is sufficient to support accurate recognition and does not necessitate changes in memory performance, (2) that mid-frontal old/new effects indexing familiarity are not merely driven by repeated access to semantic information, and (3) that priming influences neural correlates of recollection by speeding their onset. Overall, the data clearly demonstrate that there are multiple potentially interacting routes to recognition

Investigating the relationship between implicit and explicit memory: Evidence that masked repetition priming speeds the onset of recollection

Memory theories assume that unconscious processes influence conscious remembering, but the exact nature of the relationship between implicit and explicit memory remains an open question. Within the context of episodic recognition tests research typical shows that priming impacts behavioural and neural indices of familiarity. By this account, implicit memory leads to enhanced fluency of processing, which is then attributed to 'oldness' in the context of recognition judgments. Recently, however, behavioural and neuroimaging evidence has emerged to suggest that priming can also influence recollection, suggesting that the rate of recollection increases following priming. Here, we examine the relationship between priming and recollection, using Event-Related Potentials (ERPs) to assess changes in the timecourse of processing. Participants studied a series of words, and episodic memory was assessed using a standard item recognition test, but masked repetition priming preceded half of the test cues. Results confirmed that implicit memory was engaged: priming produced robust facilitation of recognition Reaction Times (RTs), with larger effects for studied than unstudied words. Mapping onto the RT data, ERPs recorded during recognition testing over centro-parietal electrodes revealed N400-like priming effects (250-500ms) that were larger in magnitude for studied than unstudied words. More importantly, priming also had a clear impact on explicit memory, as measured by recollection- related left-parietal old/new effects. While old/new effects for unprimed trials were present during the typical 500-800ms latency interval, the old/new effects seen for primed trials were equivalent in magnitude and topography, but onset ~300 ms earlier. ERPs reveal that repetition priming speeds the onset of recollection, providing a novel demonstration that unconscious memory processes can have a measureable, functional, influence on conscious remembering

Detecting the neural correlates of episodic memory with mobile EEG: Recollecting objects in the real world

Episodic memory supports recognition of the details of complex real world experiences, providing a continuous record of events embedded within spatial and temporal context. Despite the inherently dynamic nature of real events, the bulk of neuroscientific research to date examines recognition in absence of the detailed contextual information that is known to be a defining characteristic. Given the importance of environmental context for episodic memory, examining ERP correlates of memory in more naturalistic settings is vital for progress in understanding how retrieval operates in daily life. The current study capitalized on recent advances in mobile EEG technology to address this issue and is the first to investigate ERP correlates of episodic retrieval in real world contexts. Participants were guided around a pre-defined route inside a building on campus, while performing a recognition memory task, which paired images of objects with actual physical locations in the building to provide context. Importantly, the findings clearly demonstrate that it is possible to observe reliable neural correlates of memory in real world contexts. Replicating two well established ERP correlates of episodic retrieval reported in prior laboratory based studies, we detected FN400 old/new effects traditionally associated with familiarity between 300 and 500 ms, and a late posterior negativity (LPN) often linked to reconstructive processing or evaluation of retrieval outcomes between 500 and 800 ms. Moreover, the FN400 effect was found to be sensitive to retrieval of context, with more sustained effects for objects encountered in a different context at study and test. Overall, the current work highlights the power of mobile EEG technology for examining complex cognitive functions in more naturalistic real world settings

Making the case for mobile cognition: EEG and sports performance

In the high stakes world of International sport even the smallest change in performance can make the difference between success and failure, leading sports professionals to become increasingly interested in the potential benefits of neuroimaging. Here we describe evidence from EEG studies that either identify neural signals associated with expertise in sport, or employ neurofeedback to improve performance. Evidence for the validity of neurofeedback as a technique for enhancing sports performance remains limited. By contrast, progress in characterizing the neural correlates of sporting behavior is clear: frequency domain studies link expert performance to changes in alpha rhythms, whilst time-domain studies link expertise in response evaluation and motor output with modulations of P300 effects and readiness potentials. Despite early promise, however, findings have had relatively little impact for sports professionals, at least in part because there has been a mismatch between lab tasks and real sporting activity. After selectively reviewing existing findings and outlining limitations, we highlight developments in mobile EEG technology that offer new opportunities for sports neuroscience

Navigation in real-world environments : new opportunities afforded by advances in mobile brain imaging

A central question in neuroscience and psychology is how the mammalian brain represents the outside world and enables interaction with it. Significant progress on this question has been made in the domain of spatial cognition, where a consistent network of brain regions that represent external space has been identified in both humans and rodents. In rodents, much of the work to date has been done in situations where the animal is free to move about naturally. By contrast, the majority of work carried out to date in humans is static, due to limitations imposed by traditional laboratory based imaging techniques. In recent years, significant progress has been made in bridging the gap between animal and human work by employing virtual reality (VR) technology to simulate aspects of real-world navigation. Despite this progress, the VR studies often fail to fully simulate important aspects of real-world navigation, where information derived from self-motion is integrated with representations of environmental features and task goals. In the current review article, we provide a brief overview of animal and human imaging work to date, focusing on commonalties and differences in findings across species. Following on from this we discuss VR studies of spatial cognition, outlining limitations and developments, before introducing mobile brain imaging techniques and describe technical challenges and solutions for real-world recording. Finally, we discuss how these advances in mobile brain imaging technology, provide an unprecedented opportunity to illuminate how the brain represents complex multifaceted information during naturalistic navigation.Publisher PDFPeer reviewe

Navigation in Real-World Environments: New Opportunities Afforded by Advances in Mobile Brain Imaging

A central question in neuroscience and psychology is how the mammalian brain represents the outside world and enables interaction with it. Significant progress on this question has been made in the domain of spatial cognition, where a consistent network of brain regions that represent external space has been identified in both humans and rodents. In rodents, much of the work to date has been done in situations where the animal is free to move about naturally. By contrast, the majority of work carried out to date in humans is static, due to limitations imposed by traditional laboratory based imaging techniques. In recent years, significant progress has been made in bridging the gap between animal and human work by employing virtual reality (VR) technology to simulate aspects of real-world navigation. Despite this progress, the VR studies often fail to fully simulate important aspects of real-world navigation, where information derived from self-motion is integrated with representations of environmental features and task goals. In the current review article, we provide a brief overview of animal and human imaging work to date, focusing on commonalties and differences in findings across species. Following on from this we discuss VR studies of spatial cognition, outlining limitations and developments, before introducing mobile brain imaging techniques and describe technical challenges and solutions for real-world recording. Finally, we discuss how these advances in mobile brain imaging technology, provide an unprecedented opportunity to illuminate how the brain represents complex multifaceted information during naturalistic navigation

Investigating the relationship between implicit and explicit memory : evidence that masked repetition priming speeds the onset of recollection

J.L.P. was funded by the Economic and Social Research Council (www.esrc.ac.uk) through award reference ES/I901078/1. J.L.P. and D.I.D. are members of the SINAPSE collaboration (www.sinapse.ac.uk), a pooling initiative funded by the Scottish Funding Council and the Chief Scientific Office of the Scottish Executive.Memory theories assume that unconscious processes influence conscious remembering, but the exact nature of the relationship between implicit and explicit memory remains an open question. Within the context of episodic recognition tests research typical shows that priming impacts behavioral and neural indices of familiarity. By this account, implicit memory leads to enhanced fluency of processing, which is then attributed to 'oldness' in the context of recognition judgments. Recently, however, behavioral and neuroimaging evidence has emerged to suggest that priming can also influence recollection, suggesting that the rate of recollection increases following priming. Here, we examine the relationship between priming and recollection, using Event-Related Potentials (ERPs) to assess changes in the timecourse of processing. Participants studied a series of words, and episodic memory was assessed using a standard item recognition test, but masked repetition priming preceded half of the test cues. Results confirmed that implicit memory was engaged: priming produced robust facilitation of recognition Reaction Times (RTs), with larger effects for studied than unstudied words. Mapping onto the RT data, ERPs recorded during recognition testing over centro-parietal electrodes revealed N400-like priming effects (250-500 ms) that were larger in magnitude for studied than unstudied words. More importantly, priming also had a clear impact on explicit memory, as measured by recollection-related left-parietal old/new effects. While old/new effects for unprimed trials were present during the typical 500-800 ms latency interval, the old/new effects seen for primed trials were equivalent in magnitude and topography, but onset ~. 300 ms earlier. ERPs reveal that repetition priming speeds the onset of recollection, providing a novel demonstration that unconscious memory processes can have a measureable, functional, influence on conscious remembering.Publisher PDFPeer reviewe

Systematic Modification of Zingerone Reveals Structural Requirements for Attraction of Jarvis’s Fruit Fly

Tephritid fruit flies are amongst the most significant horticultural pests globally and male chemical lures are important for monitoring and control. Zingerone has emerged as a unique male fruit fly lure that can attract dacine fruit flies that are weakly or non-responsive to methyl eugenol and cuelure. However, the key features of zingerone that mediate this attraction are unknown. As Jarvis’s fruit fly, Bactrocera jarvisi (Tryon), is strongly attracted to zingerone, we evaluated the response of B. jarvisi to 37 zingerone analogues in a series of field trials to elucidate the functional groups involved in attraction. The most attractive analogues were alkoxy derivatives, with isopropoxy being the most attractive, followed by ethoxy and trifluoromethoxy analogues. All of the phenolic esters tested were also attractive with the response typically decreasing with increasing size of the ester. Results indicate that the carbonyl group, methoxy group, and phenol of zingerone are key sites for the attraction of B. jarvisi and identify some constraints on the range of structural modifications that can be made to zingerone without compromising attraction. These findings are important for future work in developing and optimising novel male chemical lures for fruit flies

Src family kinases, HCK and FGR, associate with local inflammation and tumour progression in colorectal cancer

Background: In colorectal cancer (CRC), inflammatory responses have been reported to associate with patient survival. However, the specific signalling pathways responsible for regulating inflammatory responses are not clear. Src family kinases (SFKs) impact tumourigenic processes, including inflammation. Methods: The relationship between SFK expression, inflammatory responses and cancer specific survival (CSS) in stage I-III CRC patients was assessed using immunohistochemistry on a 272 patient discovery cohort and an extended 822 patient validation cohort. Results: In the discovery cohort, cytoplasmic FGR associated with improved CSS (P=0.019), with membrane HCK (p=0.093) trending towards poorer CSS. In the validation cohort membrane FGR (p=0.016), membrane HCK (p=0.019), and cytoplasmic HCK (p=0.030) all associated with poorer CSS. Both markers also associated with decreased proliferation and cytotoxic T-lymphocytes (all p<0.05). Furthermore, cytoplasmic HCK was an independent prognostic marker compared to common clinical factors. To assess synergy a combine FGR+HCK score was assessed. The membrane FGR+HCK score strengthened associations with poor prognosis (p=0.006), decreased proliferation (p<0.001) and cytotoxic T-lymphocytes (p<0.001) Conclusions: SFKs associate with prognosis and the local inflammatory response in patients with stage I-III CRC. Active membrane FGR and HCK work in parallel to promote tumour progression and down-regulation of the local inflammatory lymphocytic response

Plasma Components Affect Accuracy of Circulating Cancer-Related MicroRNA Quantitation

Circulating microRNAs (miRNAs) have emerged as candidate biomarkers of various diseases and conditions including malignancy and pregnancy. This approach requires sensitive and accurate quantitation of miRNA concentrations in body fluids. Herein we report that enzyme-based miRNA quantitation, which is currently the mainstream approach for identifying differences in miRNA abundance among samples, is skewed by endogenous serum factors that co-purify with miRNAs and anticoagulant agents used during collection. Of importance, different miRNAs were affected to varying extent among patient samples. By developing measures to overcome these interfering activities, we increased the accuracy, and improved the sensitivity of miRNA detection up to 30-fold. Overall, the present study outlines key factors that prevent accurate miRNA quantitation in body fluids and provides approaches that enable faithful quantitation of miRNA abundance in body fluids