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Brenda Milner

Brenda Milner (born 15 July 1918) is a British-born Canadian neuropsychologist renowned as a founder of the field of neuropsychology and a pioneer in cognitive neuroscience, whose research has elucidated the neural basis of memory, language, and executive functions in the human brain.^[1]^[2] Her landmark studies, spanning over seven decades, demonstrated the functional specialization of brain regions, including the hippocampus for episodic memory and the frontal lobes for cognitive flexibility, fundamentally shaping modern understandings of brain organization and lateralization.^[2]^[3] Born in Manchester, England, to parents who were both musicians—her father a pianist and music critic, and her mother a singer—Milner displayed early aptitude in languages and mathematics rather than music.^[3] She enrolled at the University of Cambridge in 1936, initially studying mathematics before switching to psychology, and earned her M.A. in experimental psychology in 1939 amid the disruptions of World War II.^[1] Influenced by the biological psychology tradition at Cambridge and later by mentors such as Donald Hebb, she pursued doctoral studies at McGill University in Montreal, completing her Ph.D. in 1952 on the effects of temporal lobe damage on cognition.^[1]^[3] In 1950, Milner joined the Montreal Neurological Institute (The Neuro) at McGill, where she collaborated with neurosurgeon Wilder Penfield on patients undergoing epilepsy surgery, marking the beginning of her lifelong career there, now exceeding 70 years.^[1] Her seminal work with patient H.M. (Henry Molaison), beginning in 1955 and continuing for over three decades, revealed the critical role of the hippocampus in forming new declarative memories while sparing other cognitive abilities, leading to the identification of distinct short-term and long-term memory systems.^[2] She further advanced knowledge of frontal lobe functions through systematic assessments using tools like the Wisconsin Card Sorting Test, uncovering deficits in perseveration and abstract reasoning following dorsolateral prefrontal excisions, and highlighting hemispheric asymmetries in cognitive processing.^[2] Milner's research extended to bilingualism, spatial memory, and the effects of aging on cognition, often integrating lesion studies with emerging neuroimaging techniques in her later work.^[1]^[3] Throughout her career, Milner has received numerous prestigious accolades, including the Kavli Prize in Neuroscience (2014) shared for advancing understanding of neural basis of cognition, the Balzan Prize in Cognitive Neuroscience (2009), the Gairdner International Award (2005), and the National Academy of Sciences Award in the Neurosciences (2004).^[4] More recently, she was inducted into Canada's Walk of Fame (2023) and received the Prix Grands Sages du scientifique en chef from the Quebec government (2023), recognizing her enduring impact.^[4] As Professor Emerita in the Department of Neurology and Neurosurgery at McGill and a member of The Neuro, she continues to influence the field at age 107.^[1]

Early life and education

Early life

Brenda Milner was born on July 15, 1918, in Manchester, England, to parents immersed in the world of music. Her father, Samuel Langford, served as the music critic for the Manchester Guardian and was also a pianist, while her mother, Leslie Doig, was a singer who had studied under him. The family home was filled with intellectual stimulation, including discussions of literature and the arts, though Milner survived the 1918 influenza pandemic alongside her mother in its aftermath.^[5]^[6] From an early age, Milner displayed no talent for music despite her parents' influence, instead gravitating toward scientific and analytical pursuits. She enjoyed literature, languages, mathematics, physics, and solving puzzles, which highlighted her logical mindset within the family's encouraging intellectual environment. Her father provided homeschooling in classics until his death from tuberculosis when she was eight, after which her mother taught her French, fostering a lifelong appreciation for languages in which she became fluent in English, French, Latin, Italian, and some German.^[3]^[5]^[6] At age eight, Milner enrolled at Withington Girls’ School in Manchester, where she thrived in languages and developed a strong passion for mathematics during the interwar period—a time of economic uncertainty and cultural shifts in England following World War I. Her early years there were marked by academic focus amid the looming tensions that would soon erupt into World War II, which disrupted broader societal life and her emerging youth by 1939 through air raid precautions and national mobilization efforts.^[5]^[6] These formative experiences shaped her initial inclinations toward mathematics and the sciences, setting the stage for her later transition to studies in experimental psychology.^[3]^[5]

Education

Milner enrolled at Newnham College, University of Cambridge, in 1936, initially intending to study mathematics before switching to experimental psychology.30520-7) She graduated with a B.A. in experimental psychology in 1939, a degree then classified under moral sciences.^[7] Following her undergraduate degree, she held a two-year Sarah Smithson Research Studentship at Newnham College from 1939 to 1941, conducting perceptual research in the Psychological Laboratory under the supervision of Frederic C. Bartlett, which included early experiments on visual perception and learning.^[8] The outbreak of World War II interrupted her postgraduate work at Cambridge, as laboratory efforts shifted toward applied psychological testing for air crew selection, incorporating assessments of visual and perceptual abilities.^[9] She later completed her M.A. at Cambridge in 1949.^[7] In 1944, Milner relocated to Montreal, Canada, with her husband, Peter Milner, an electrical engineer recruited for wartime nuclear research.^[10] Upon arriving in Canada, Milner taught psychology courses at the Université de Montréal, focusing on experimental methods, while preparing for advanced graduate study.^[11] In 1950, she enrolled in the Ph.D. program in physiological psychology at McGill University under the supervision of Donald O. Hebb, whose theories on neural organization and behavior profoundly shaped her approach.^[3] Her doctoral coursework emphasized the physiological bases of learning and cognition, including seminars on brain function and behavioral deficits.^[8] Early laboratory experiences at McGill involved testing patients with neurological lesions in Hebb's setup, fostering her interest in linking psychological processes to brain structures.^[12] Milner's Ph.D. thesis, titled Intellectual Effects of Temporal-Lobe Damage in Man, examined cognitive impairments following temporal lobe excisions, drawing on clinical data from neurosurgical cases; she defended it successfully in 1952.^[13] Concurrently, starting in 1950, she collaborated at the Montreal Neurological Institute under Wilder Penfield, whose epilepsy surgery patients provided critical opportunities to observe and assess brain-behavior relationships, complementing Hebb's theoretical framework.^[3]

Professional career

Early positions

Following her undergraduate degree, Milner served as a research assistant at the University of Cambridge from 1939 to 1942, where her postgraduate work was redirected toward the Allied war effort with the outbreak of World War II. She contributed to a team evaluating perceptual tasks for selecting Royal Air Force (RAF) aircrew, specifically studying visual tracking abilities and fatigue to differentiate suitable candidates for fighter pilots versus bomber pilots.^[14]^[9] In 1944, Milner relocated to Montreal, Canada, with her husband, Peter Milner, who had accepted a position in atomic research. She immediately took up a role as Professeur agrégé at the Institut de Psychologie of the Université de Montréal, serving from 1944 to 1952 and teaching courses in experimental psychology, including animal behavior and learning. During this time, she attended psychology seminars at nearby McGill University, where she first encountered Donald O. Hebb, the department chair, whose ideas on brain organization profoundly influenced her.^[1]^[3]^[15] From 1950 to 1952, while completing her PhD under Hebb's supervision at McGill—briefly referencing her training in physiological psychology—Milner held a research associate position in the Psychology Department, concentrating on animal behavior and the effects of brain lesions on learning. Her work involved experimental studies with rats, examining how lesions disrupted performance in tasks like maze navigation, providing early insights into neural substrates of cognition.^[15]^[3] Concurrently, Milner gained initial clinical exposure at the Montreal Neurological Institute (MNI) through Wilder Penfield's epilepsy surgery program, beginning around 1950 as part of her doctoral research. She assisted in developing standardized testing protocols in the early 1950s to evaluate patients' cognitive abilities pre- and post-operatively, focusing on memory and perceptual functions to inform surgical decisions. This hands-on experience bridged her animal studies with human neuropsychology.^[3]^[1] These early roles culminated in her first publications on brain-behavior relations in the 1950s, including papers detailing lesion effects on maze learning in rats, which highlighted regional brain contributions to adaptive behavior and laid groundwork for her later human studies. For instance, her analyses showed that certain temporal lobe lesions impaired spatial navigation without affecting basic motor skills.^[3]

Work at McGill University and Montreal Neurological Institute

Brenda Milner joined the Montreal Neurological Institute (MNI) in 1950, initially supported by a grant from Donald Hebb, and began her formal academic career at McGill University as a research associate in 1952, the year she completed her PhD there. She was appointed lecturer in the Department of Psychology from 1953 to 1960, followed by promotion to assistant professor from 1960 to 1964 and associate professor from 1964 to 1970, before rising to full professor in the Department of Neurology and Neurosurgery in 1970, later becoming Professor Emerita, alongside her role as Dorothy J. Killam Professor at the MNI.^[8]^[1] Milner's early work at the MNI involved close collaboration with Wilder Penfield, the institute's founder and director, on patients undergoing surgery for epilepsy. Starting in 1950, she conducted pre- and post-operative assessments of these patients, which necessitated the development of standardized neuropsychological testing batteries to systematically evaluate cognitive functions affected by brain lesions. These batteries, refined through her clinical observations and published in key works such as her 1954 review in Psychological Bulletin and a 1958 co-authored paper with Penfield in A.M.A. Archives of Neurology and Psychiatry, formed the foundation for rigorous, repeatable assessments in neuropsychology at the MNI.^[7]^[8]^[12] In 1972, Milner assumed leadership of the Neuropsychology Unit at the MNI, which she helped evolve into the Cognitive Neuroscience Unit by the 1980s, fostering an interdisciplinary environment that integrated behavioral testing with emerging neuroimaging techniques. Under her direction, which continued until 1990 when she handed over day-to-day operations to Michael Petrides, the unit trained generations of researchers, including over a dozen PhD students such as Doreen Kimura, Marilyn Jones-Gotman, and Philip Corsi, who advanced the field through their subsequent contributions.^[1]^[7] Milner's institutional efforts extended to establishing dedicated laboratory facilities at the MNI for lesion studies in the mid-1960s, supported by Medical Research Council grants that enabled expanded testing capabilities beyond initial weekend sessions. By 1989, she co-founded the McConnell Brain Imaging Centre with Petrides using a McDonnell-Pew grant, facilitating the integration of positron emission tomography (PET) and magnetic resonance imaging (MRI) into cognitive research protocols, with active lesion and imaging studies continuing into the 2000s. These developments solidified the MNI as a global hub for cognitive neuroscience, emphasizing collaborative, patient-centered methodologies.^[7]^[1]^[8]

Research contributions

Memory systems and the hippocampus

Brenda Milner's pioneering studies on patient H.M., conducted in collaboration with neurosurgeon William Beecher Scoville and later with neuropsychologist Suzanne Corkin during the 1950s and 1960s, provided critical evidence for the role of the hippocampus in memory formation.^[16] In 1953, Scoville performed a bilateral resection of the medial temporal lobes, including the hippocampus, on H.M. to alleviate severe epilepsy, resulting in profound anterograde amnesia that prevented the formation of new episodic memories while leaving remote memories and other cognitive functions largely intact.^[17] Milner's detailed neuropsychological assessments of H.M. revealed that this surgery specifically disrupted the ability to consolidate new declarative memories—those involving facts and events—while sparing other memory processes.^[18] In their seminal 1957 paper, Scoville and Milner documented H.M.'s preserved intellectual abilities, such as normal IQ scores and intact short-term verbal memory (e.g., digit span of seven forward and five backward), alongside severe impairment in long-term episodic memory, where H.M. could not recall events from minutes earlier without cues.^[17] This dissociation highlighted that short-term memory storage, limited to immediate recall, remained functional post-surgery, but the transfer to long-term storage for conscious recollection was abolished.^[18] These findings challenged unitary theories of memory and laid the groundwork for understanding the hippocampus's selective involvement in declarative memory systems.^[16] Building on these observations, Milner's work with H.M. and Corkin established the concept of multiple dissociable memory systems in the brain.^[3] Declarative memory, which is hippocampus-dependent and supports explicit recall of personal experiences and factual knowledge, was severely impaired in H.M., whereas procedural memory—encompassing skill-based learning such as motor habits—was hippocampus-independent and remained intact.^[16] This framework demonstrated that procedural memory operates implicitly, without requiring conscious awareness or hippocampal mediation, allowing amnesic patients to acquire new abilities over repeated exposure.^[19] A key experimental paradigm illustrating this dissociation was the mirror-tracing task, designed by Milner to assess procedural learning.^[3] In this task, H.M. was required to trace the outline of a five-pointed star viewed only in a mirror, which reverses visual feedback and demands hand-eye coordination adaptation. Despite no recollection of prior sessions, H.M. showed progressive improvement across trials, reducing errors and time taken, comparable to healthy controls—evidence of intact procedural memory acquisition without episodic encoding.^[16] Such paradigms underscored the modular organization of memory, with the hippocampus gating declarative but not procedural systems, influencing subsequent models of human cognition.^[19]

Hemispheric specialization

Brenda Milner's research on hemispheric specialization, conducted primarily in the 1960s and 1970s at the Montreal Neurological Institute, utilized patients with unilateral brain lesions and those who had undergone cerebral commissurotomy to delineate functional asymmetries between the left and right hemispheres.^[20] Her studies with patients having unilateral temporal or frontal lobe excisions revealed that left-hemisphere damage often impaired language-related tasks and sequential processing, such as verbal fluency and serial ordering, while right-hemisphere lesions disrupted visuospatial abilities, including maze navigation and design fluency.^[12] For instance, left frontal resections led to significant deficits in verbal recency judgments and articulation of sequences, underscoring the left hemisphere's dominance in analytical and linguistic functions.^[21] In parallel, Milner's investigations of split-brain patients, who had undergone sectioning of the corpus callosum to treat epilepsy, provided critical evidence for interhemispheric independence. Testing these individuals on tactile and visual matching tasks demonstrated the right hemisphere's superior performance in processing unfamiliar patterns and shapes, particularly when stimuli were presented to the left hand or visual field, which projects contralaterally.^[22] A seminal 1972 publication by Milner and Jeeves on tactile pattern recognition, along with her 1974 synthesis, highlighted asymmetries in tactile-visual integration and argued for specialized localization of psychological processes, with the left hemisphere excelling in verbal and propositional tasks, and the right in nonverbal, holistic ones. This work built on earlier observations, such as her 1968 study showing that right temporal-lobe excisions impaired visual recognition and recall of faces, supporting the right hemisphere's role in facial processing.^[23] Milner's contributions extended to language lateralization, where she employed dichotic listening tests and the intracarotid sodium amobarbital (Wada) procedure to map dominance, revealing that most right-handers exhibit left-hemisphere control of speech production in Broca's area and comprehension in Wernicke's area.^[20] These methods quantified atypical lateralization in some individuals, influencing understandings of how hemispheric asymmetries underpin cognitive modularity. Her patient-based approach, overlapping briefly with memory assessments in lesioned cases, established enduring frameworks for studying brain lateralization without relying on invasive imaging.

Bilingualism and spatial processing

During the 1990s and 2000s, Brenda Milner conducted studies on healthy bilingual participants that demonstrated activation patterns in frontal lobe regions during language tasks, building on her earlier work on hemispheric lateralization. In positron emission tomography (PET) experiments, bilingual participants showed activation in the left frontal cortex when generating words in their first and second languages, suggesting common neural substrates for within- and across-language processing.^[24] In collaboration with Denise Klein, Milner utilized PET imaging to map bilingual activation patterns, revealing distinct yet overlapping neural networks for first and second languages. Their research on English-French and Chinese-English bilinguals identified involvement of the left putamen and prefrontal areas during second-language production. For instance, verb generation tasks across languages elicited frontal activations, underscoring the role of executive control in language processing.^[25] This work established that bilingualism engages frontal networks to support language use.^[26] Milner's investigations into spatial processing employed tasks such as object-location recall to probe navigation abilities, linking right hippocampal and parietal cortex activations to spatial orientation and memory. In studies of patients with medial temporal lobe lesions, right hippocampal damage impaired recall of object locations and route learning, while parietal regions supported visuospatial integration for effective navigation. These experiments demonstrated that the right hippocampus encodes spatial layouts, with parietal inputs providing egocentric reference frames essential for wayfinding.^[27]^[28] At the Montreal Neurological Institute (MNI), Milner's research extended to the effects of aging on cognition, often integrating lesion studies with emerging neuroimaging techniques.^[1]

Later career and legacy

Recent research activities

Since the early 2010s, Milner has employed functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) to delineate memory networks in healthy aging adults, offering neuroimaging validation for her foundational lesion studies on hippocampal and frontal contributions to memory.^[29]^[30] In the 2020s, her research has emphasized cognitive resilience in advanced age, with collaborations involving postdoctoral mentees to explore interhemispheric cooperation in memory retrieval and the supportive role of visual processing in abstract memory formation—efforts aimed at informing interventions for dementia, brain injury, and learning impairments.^[31] A 2023 interview captured Milner's personal reflections on sustained cognitive function at age 104, illustrating principles of resilience observed in centenarians through her longitudinal perspective on memory preservation.^[32] The 2025 BrainFacts.org profile recognizes her persistent engagement with memory mechanisms.^[33] In 2025, on her 107th birthday, Milner continued her research interests in hemispheric cooperation for memory, as highlighted in profiles celebrating her enduring contributions.^[31] Milner has investigated bilingualism using neuroimaging, including studies on language pathways and the effects of aging on cognition.^[9]

Influence on neuropsychology

Brenda Milner's pioneering research fundamentally shaped neuropsychology by establishing it as a rigorous, empirically grounded discipline focused on linking specific brain regions to cognitive functions, moving beyond earlier simplistic localization theories toward a modular understanding of brain organization. Her seminal 1957 collaboration with William Beecher Scoville on patient H.M. demonstrated the medial temporal lobes' critical role in episodic memory formation while sparing other memory types, such as procedural learning, thereby introducing the concept of dissociable memory systems that revolutionized how researchers conceptualize brain modularity.^[17] This paradigm shift influenced subsequent models in cognitive neuroscience, emphasizing specialized neural circuits for distinct cognitive processes. Widely recognized as a founder of neuropsychology, Milner's integration of clinical observation with behavioral testing at the Montreal Neurological Institute (MNI) transformed the field from anecdotal case studies to systematic investigation, solidifying her legacy as the discipline's architect.^[34]^[35] Through her mentorship, Milner trained numerous graduate students and postdoctoral fellows who emerged as leaders in cognitive neuroscience, fostering a global network that extended her methodologies worldwide. Notable trainees include Doreen Kimura, who advanced studies on hemispheric lateralization; Marilyn Jones-Gotman, a key figure in olfactory and gustatory research; and Michael Petrides, who contributed to prefrontal cortex mapping, with many establishing prominent programs at institutions like the University of Western Ontario and McGill University.^[7] She supervised over a dozen PhD students and emphasized hands-on clinical training, ensuring her protégés combined patient-based insights with experimental rigor, which amplified neuropsychology's reach into academic and clinical settings globally.^[1] In 2007, she furthered this influence by founding the Brenda Milner Foundation to support postdoctoral fellowships in cognitive neuroscience at the MNI, perpetuating her emphasis on interdisciplinary brain research.^[1] Milner's methodological innovations, particularly the adaptation of the intracarotid amobarbital (Wada) procedure for preoperative memory assessment, provided a standardized tool to evaluate hemispheric contributions to cognition, remaining a cornerstone in epilepsy surgery protocols today.^[36] She also adapted the Wisconsin Card Sorting Test to probe frontal lobe executive functions, enabling precise quantification of cognitive deficits post-surgery and influencing diagnostic practices in clinical neuropsychology.^[12] These tools, alongside her detailed lesion-behavior correlations, elevated MNI to a preeminent international hub for neuropsychology, drawing collaborators like Roger Sperry and Ennio De Renzi and establishing Montreal as a epicenter for brain-behavior studies that continue to inform global research paradigms.^[7]

Awards and honors

Major scientific prizes

Brenda Milner has received several prestigious international scientific prizes recognizing her foundational contributions to cognitive neuroscience, particularly in understanding memory systems and brain lateralization. These awards highlight her pioneering patient-based studies that delineated the roles of specific brain regions in cognition. In 1973, Milner was awarded the American Psychological Association's Distinguished Scientific Contribution Award for her seminal work on the neuropsychology of memory, including analyses of patients with temporal lobe lesions that revealed dissociations between different memory types, such as episodic and procedural memory.^[37] This honor underscored her early collaborations with Wilder Penfield at the Montreal Neurological Institute, where her research established key frameworks for integrating behavioral data with neuroanatomical findings.^[4] The 1987 Ralph W. Gerard Prize from the Society for Neuroscience acknowledged Milner's groundbreaking investigations into hemispheric specialization and memory processing, notably her studies demonstrating the distinct functions of the left and right temporal lobes in verbal and nonverbal memory tasks. This prize, one of the field's highest honors for lifetime achievement in neuroscience, celebrated her rigorous experimental approaches that advanced the understanding of brain asymmetry in cognitive functions.^[4] In 2004, Milner received the National Academy of Sciences Award in the Neurosciences for her discoveries concerning the neural substrates of memory and cognition.^[4] In 2005, Milner received the Canada Gairdner International Award for her pioneering research on human memory mechanisms, providing an essential framework for correlating neuroanatomical, neurophysiological, and neuropharmacological insights into hippocampal and frontal lobe functions.^[38] Her work, initiated in the 1950s with patient H.M., illuminated the hippocampus's critical role in forming new declarative memories while sparing other cognitive abilities.^[38] The 2009 Balzan Prize in Cognitive Neurosciences was bestowed upon Milner for her pioneering studies on the hippocampus's role in memory formation and the identification of distinct memory systems, including her demonstrations of preserved skill learning in amnesic patients.^[39] Valued at one million Swiss francs, the award emphasized her influence on research into brain regions like the frontal cortex and the implications for disorders such as Alzheimer's disease.^[39] In 2011, she was awarded the Pearl Meister Greengard Prize from Rockefeller University for her contributions to neuroscience.^[4] In 2014, Milner shared the Kavli Prize in Neuroscience with John O'Keefe and Marcus E. Raichle for the discovery of specialized brain networks underlying memory and cognition, with her contributions focusing on the medial temporal lobes' orchestration of episodic memory.^[40] She also received the Dan David Prize that year for her work on the human brain.^[4] The $1 million prize, administered by the Norwegian Academy of Science and Letters, recognized how her findings complemented O'Keefe's spatial mapping work and Raichle's brain imaging advancements to map cognitive processes.^[40]

National and honorary recognitions

Brenda Milner was appointed an Officer of the Order of Canada in 1984 and promoted to Companion, the highest level of the order, in 2004 for her pioneering contributions to neuropsychology.^[41]^[9] She also received the Grand Officer distinction in the Order of Quebec in 2009, recognizing her impact on Quebec's scientific community.^[29] Milner was elected a Fellow of the Royal Society of Canada in 1976, acknowledging her early advancements in cognitive neuroscience.^[9]^[3] In 1979, she became a Fellow of the Royal Society (UK) for her discoveries on specialized brain networks for memory and cognition.^[42] She was named a Foreign Honorary Member of the American Academy of Arts and Sciences in 2005, honoring her international influence in behavioral neuroscience.^[43]^[9] In 2023, she received the Prix Grands Sages du scientifique en chef from the Quebec government and was inducted into Canada's Walk of Fame, recognizing her enduring contributions to science.^[4] Throughout her career, Milner has been awarded more than 25 honorary doctorates from universities in Canada, the United States, and Europe, reflecting her enduring academic stature as of 2020.^[5] Notable examples include a Doctor of Laws from McGill University in 1991, where she has spent much of her professional life, and a Doctor of Science from the University of Cambridge in 2000, her alma mater.^[44]^[45] In 2025, Milner's legacy continued to be highlighted in profiles celebrating women in STEM, such as features on her role as a trailblazer in neuropsychology for the International Day of Women and Girls in Science and Women's History Month initiatives.^[46]^[47]

References

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[41]
Brenda Milner, CC, GOQ, DSc, PhD - McGill University
Dr. Brenda Milner is the Dorothy J. Killam Professor at The Neuro, and a professor in the Department of Neurology and Neurosurgery, McGill University.
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Dr. Brenda Milner
She set the tone of her career very early on by going against Hebb's advice to study patients of Dr. Wilder Penfield at the Montreal Neurological Institute.<|separator|>
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Selected Honorands - Honorary degrees - University of Cambridge
DegreeDoctor of Law, Year2000. NameEdward Alan John George, DegreeDoctor of Law, Year2000. NameBrenda Milner, DegreeDoctor of Law*, Year2000. NameElias James ...
[44]
From Struggle to Success: The Legacy of Women in Science
Feb 11, 2025 · WiNUK Awards 2025 ... Brenda Milner. Image taken from McGill.ca. Brenda is considered to be a founder of clinical neuropsychology and cognitive ...
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Dr. Brenda Milner: A Pioneer in Neuropsychology - LinkedIn
Oct 9, 2025 · Her studies using brain imaging and surgery continue to deepen our understanding of the hippocampus and its crucial role in memory formation.Missing: setup lesion MNI