Warren H. Meck

The Timing Research Forum is dedicating this page to Warren Meck. Without Warren’s vision, kindness, and stewardship of the timing field and community, neither would TRF exist nor would there be as many researchers working on time as we’re lucky to have amongst us today.

This webpage is one of a series of actions that TRF is taking for honoring, celebrating, and remembering Warren. Please join us in these efforts.

Below this obituary we encourage all of you to share your memories of Warren. You can send your text (and pictures if you like) to TRF’s email (trf@timingforum.org) and we will post your contribution on this page.

Warren H. Meck

November 17, 1956 – January 21, 2020

Warren Meck’s profound impact on the field of timing and time perception spanned four decades. From the 1980s through the 2010s, he and his collaborators consistently produced seminal papers that advanced our fundamental understanding of timing in the seconds to minutes range (i.e., interval timing). Warren played a crucial role in the development of two dominant theories of interval timing, Scalar Timing Theory (Gibbon, Church, & Meck, 1984) and the Striatal Beat-Frequency Model (Matell & Meck, 2000). His research was marked by creativity, independence of thought, and an openness to consider and incorporate ideas from other domains. Moreover, although most of his empirical research relied on rodent models, he produced a substantial body of research using human participants. Consequently, Warren facilitated communication and transmission of ideas among researchers using animal models and those using humans, resulting in a richer conceptualization of the cognitive and neural mechanisms of interval timing.

Warren’s dedication to his fellow researchers, whether students or colleagues, was unequalled. Most members of the Timing Research Forum are aware of the book and special journal issues he organized, as well as the critical role he played in founding Timing & Time Perception. Many will have discussed their own research with him and found him to be highly engaged with and encouraging of their work. Few will know that he once checked himself out of hospital, against doctor’s advice, because a collaborator was depending on him to make an invited presentation.

Warren grew up on a farm in eastern Pennsylvania and as a teenager there were glimpses of the independence that marked his academic career. Indeed, he may have been the first high-school football player in Pennsylvania history to give up a coveted team position to join the school marching band. Following high school, Warren spent two years at Pennsylvania State University before enrolling at the University of California – San Diego to complete a BA in Psychology. At UCSD, Warren worked in Ed Fantino’s lab, which resulted in his first academic publication (Fantino, Dunn, & Meck, 1979).

In 1978, Warren moved to Brown University to pursue a PhD in psychology under the mentorship of Russell Church. As the story goes, he decided to drive from California to Rhode Island by routing through Canada. Intriguingly, he was temporarily stopped at the border as he tried to re-enter the United States because his wallet was, allegedly, found at the site where a Navy helicopter disappeared. One might imagine how time stood still as he was questioned by the border patrol…perhaps foretelling his developing interest in temporal relativity. His PhD, for which he won the James McKeen Cattell Dissertation Award, was awarded in 1982. He remained at Brown for another four years, serving first as a research scientist and then as an Assistant Professor.

In 1986, Warren joined Columbia University as an Assistant Professor and was promoted to Associate Professor in 1990. In 1994, the year he received the Early Career Recognition Award from the Eastern Psychological Association, he moved to the Psychology Department at Duke University. He was promoted to full professor at Duke in 2001 and received the James McKeen Cattell Sabbatical Fellowship shortly thereafter. Throughout his career, Warren’s research was supported by grants from several National Institute of Health agencies, as well as the National Science Foundation, and the Welcome Trust, among others.

A journey through (interval) timing with Warren H. Meck

Brown University: 1978 – 1986
At Brown, Warren conducted a mixture of purely behavioral research, such as examining cross-modal transfer of temporal rules (e.g., Meck & Church, 1982a, 1982b), and research that combined behavioral measures with pharmacological or brain lesion manipulations (e.g., Meck, 1983; Meck, Church, & Olton, 1984). He joined Russ Church in a collaboration with John Gibbon that resulted in the development of Scalar Timing Theory (STT), the information processing model based on Scalar Expectancy Theory, and refinement of its mathematical instantiation (Gibbon, Church, & Meck, 1984). Over the next 15 years, STT became the dominant interpretive framework for studies of timing in both human and non-human animals.

Even to investigators far outside the field, Warren was instantly recognizable as a major pioneer in revealing the neural mechanisms of interval timing. His pharmacological work demonstrated distinct effects of dopaminergic and cholinergic drugs on timing. Specifically, manipulation of the dopaminergic system primarily affected the clock stage within the STT framework, whereas manipulation of the cholinergic system primarily affected the memory stage (e.g., Meck, 1983, 1986, 1987; Meck, Church, & Wenk, 1986; Meck & Church, 1987). At Brown, Warren began to make the connections between putative cognitive timing mechanisms and underlying neural substrates which defined his career.

During this period, Warren and Russ initiated a very successful collaboration with David Olton. Indeed, their 1984 Behavioural Neuroscience paper, entitled Hippocampus, Time and Memory (Meck, Church, & Olton, 1984) was recognized by the journal as a “classic” publication on the occasion of its 30th anniversary celebration. The paper showed that fimbria-fornix (FFx) lesions distorted long-term temporal memory and impaired short-term memory only when there were gaps in the timing signal, thereby distinguishing mechanisms of duration perception and temporal memory. Importantly, the paper encouraged “the development of computational models with plausible neural mechanisms …, the use of multiple behavioral measures of timing, and empirical research on the neural mechanisms of timing and temporal memory using ensemble recording of neurons in prefrontal-striatal-hippocampal circuits” (Meck, Church, & Matell, 2013).

Dissecting the internal clock using careful behavioral, pharmacological and lesion manipulations, a hallmark of Warren’s research, started at Brown, and continued throughout his career. It was also at Brown that students and colleagues became aware of his extreme night owl tendencies, which were possibly the source of the rumors that he, literally, lived in his office.

Columbia University: 1986 – 1994
Warren met Christina Williams, who was a faculty member in the Psychology Department of Barnard College, when he joined Columbia. This collaboration resulted in influential work on the effects of hormones and maternal diet on the development of spatial and temporal learning and memory (e.g., Williams, Barnett, & Meck 1990; Williams & Meck, 1991). In addition to becoming scientific partners, Warren and Tina also became marriage partners, with Tina becoming jointly responsible for Pilot, an extraordinary Shih Tzu. Pilot was the first in a line of Shih Tzus ruling over the Meck-Williams household, succeeded by the exuberant Million Williams, and with the position currently held by the extremely rascally Riley.

Warren continued to publish lesion and pharmacological studies of interval timing throughout his years at Columbia (e.g., Meck et al., 1987; Meck, 1988), as well as granular analyses of timing behaviour, including a seminal single-trials analysis of peak-interval procedure performance in rats, which supported a parallel scalar timing model rather than a serial or quasi-serial timing model (Church, Meck, & Gibbon, 1994).

Warren’s research relationship with John Gibbon also deepened during this period. Warren and his students at the time (Nancy Dallal, Sean Hinton, Trevor Penney, Brian Rakitin) regularly made the trip from the main campus at 116th Street up to John’s lab at 168th Street for John’s Friday lunchtime lab meeting, which provided opportunities for significant cross-fertilization of ideas. One consequence was a new line of research with human participants that used tasks originally developed for non-human animals (i.e., duration bisection, peak interval procedure). Around the same time, Chara Malapani, from the Hôpital de la Salpêtrière in Paris, initiated a collaboration with Warren and John to study timing in patients with Parkinson’s disease.

Warren’s reputation as a highly dedicated and caring graduate supervisor started at Columbia. He was available for conversations about research, and life, whenever needed. On at least one Thanksgiving holiday, he brought a student, unannounced, for a weekend visit to Tina’s parents. The student was grateful not to be alone at Thanksgiving and very happy to be a junior member of the “Timing Mafia”.

Duke University: 1994 – 2020
Although Warren’s main focus remained non-human animal research throughout the late 90’s and early 00’s, he continued to pursue human timing research both with students and colleagues at Duke and with collaborators elsewhere, including Viviane Pouthas and Sylvie Droit-Volet in France. Indeed, most of the human timing research initiated at Columbia University was published after Warren joined Duke. These publications included application of the peak interval procedure and single trial analysis to college students, which demonstrated qualitatively equivalent performance between humans and rodents on the peak-interval procedure (Rakitin et al., 1998), an on-off dopamine supplementation study in patients with Parkinson’s Disease which revealed a coupling of temporal memories in the off state (Malapani et al., 1998), and a study of stimulus modality effects in the duration bisection task which addressed the question of why “sounds are longer than lights” (Penney, Gibbon, & Meck, 2000). These three papers, among others, extended and expanded work that Warren had started with rodents in the 80’s.

Warren and Tina continued their close scientific collaboration, in addition to their service to the Shih Tzus, throughout their years at Duke. Under Tina’s influence, Warren further developed his interest in time’s close cousin – space – and how they work together in memory (e.g., Meck & Williams 1997a, 1997b). This collaboration also helped inspire a perspective on neurobiological mechanisms that went beyond ‘pushing the system’ with lesions or short-term drug administration to understanding more naturalistic influences that affect normal variation. These included the effects of sex hormones on timing, spatial memory, and other abilities, and how pre-natal choline supplementation could affect cognition throughout the lifespan and even the trajectory of aging (Meck & Williams, 2003; Williams, Meck, Heyer, & Loy, 1998). The projects with Tina were also influential in Warren’s increasing incorporation of advanced neuroscience methods to better understand the changes in neural structure and function underlying these effects (e.g., Agostino, Cheng, Williams, West, & Meck, 2013; Cheng, Meck, & Williams, 2006; Mellott, Williams, & Meck, 2004).

Human timing projects developed at Duke also began to appear in print, such as Sean Hinton’s work investigating nicotine effects on timing (Hinton & Meck, 1996), Cindy Lustig’s examination of the effects of time of day and aging on duration bisection performance (Lustig & Meck, 2001), and Elizabeth Brannon’s event related potential investigations of timing in infants (e.g., Brannon, Roussel, Meck, & Woldorff, 2004).

The work with animal models continued apace, resulting in new techniques and, subsequently, significant theoretical advances. For example, Matthew Matell and Warren developed the bi- and tri-peak versions of the peak-interval procedure (e.g., Matell & Meck, 1999), which were later used to refine our understanding of the role of sensitization in drug-induced alterations of time perception (Matell, King, & Meck, 2004) and the role of cortical and striatal neurons in timing (e.g., Matell, Meck, & Nicolelis, 2003). Meanwhile, he and Catalin Buhusi developed the reverse-gap procedure (Buhusi & Meck, 2000) and, in one of many papers using this protocol, demonstrated that dopaminergic drugs induce clock and attentional effects (Buhusi & Meck, 2002), rather than merely clock effects, as had been suggested by Warren’s earlier work.

One of the marks of Warren’s rigor and innovation as a scientist was his willingness to leave behind the ideas and models on which he built his reputation for new ideas, constructed with his students, that were more informed by recent developments in neurobiology as well as the new behavioural probes developed in his own lab. The most important contribution was the Striatal Beat-Frequency (SBF) model, which he and Matthew Matell first proposed in a paper published in 2000 (Matell & Meck, 2000). The SBF model grew out of STT in the sense that behavioral and neuroscience research inspired and interpreted within the STT framework provided the foundational material that the SBF model built upon, but SBF conceptualized a physiologically viable solution to the question of how brains time. Two highly cited review papers, which clarify the shortcomings of STT and explain how the SBF model overcomes those shortcomings, were subsequently published (Matell & Meck, 2004; Buhusi & Meck, 2005). Warren, Matt, and their collaborator Miguel Nicolelis, also provided the first direct physiological support for the model via a series of ensemble recording experiments from the striatum and cortical regions in the rat (Matell, Meck, & Nicolelis, 2003).

Both the model and those papers were undoubtedly shaped by engaging discussions lasting well into the night during his graduate seminar on timing and time perception. Consistent with his generosity and night-owl tendencies, Warren “made a deal” with the students that class meetings would take place in the evenings at his Renaissance Revival villa (a short walk from campus), in front of a roaring fire, and accompanied by pizza and beer. No doubt fuelled by those refreshments, as well as a passion for ideas, Warren and his graduate students were very productive in the period from 2004 to 2009. Ruey-Kuang Cheng first-authored 10 articles, mainly sophisticated pharmacological dissections of the timing circuit (e.g., Cheng, Meck, & Williams, 2006; Cheng, MacDonald & Meck, 2006) and Chris MacDonald first-authored five articles, including important papers on the relationship between interval timing and reaction time (e.g., MacDonald & Meck, 2006).

Projects developed with collaborators, some of whom were former students, also came to fruition during this “peak interval” in publication output. Warren and Catalin Buhusi continued to work very productively together with multiple publications using the gap and reverse-gap procedures. Consistent with Warren’s efforts to incorporate (and infiltrate) other fields, a paper with Cindy Lustig and Matt Matell (Lustig, Matell, & Meck, 2005) examined how theories of working memory based largely on human studies could be integrated with Matt’s SBF model of cortical-striatal interactions to explain how the same neural representations could support both identity and duration information. Matt Matell and Melissa Bateson applied a single-trial analysis to peak-interval data obtained while rats were on or off methamphetamine to show leftward shifts in peak functions were due to clock speed effects (Matell, Bateson, & Meck, 2006).

Several papers that had extremely long gestation periods were also published in this period. One, a multi-species study that showed similar timing-errors in humans, pigeons and mice, thereby suggesting a common mechanism, used data collected over a 20-year span (Penney, Gibbon, & Meck, 2008). A draft of another paper existed for close to 20 years prior to finally being published (Meck, 2006). Both publications reflected Warren’s habit to continue thinking about and working with old(er) data, which, like wine, could improve with age.

This tendency to think about old data and research themes manifested in a continual effort to refine and improve old theories by applying new techniques and analysis approaches to the questions those theories addressed. Hence, in the 2010s, many of Warren’s publications revisited themes from earlier in his career, but from new directions and with new techniques. Much of this work was pursued with students, some of whom were Warren’s advisee’s (Bon-Mi Gu, Bin Yin, Elijah Petter, and Nicholas Lusk), whereas others (e.g., Jessica Lake, Sundeep Teki, Tadeusz Kononowicz) were “on loan” from other labs to pursue collaborative projects with their own advisors (e.g., Kevin LaBar, Tim Griffiths, Hedderik van Rijn) and Warren.

Warren, Bon-Mi Gu, and Hedderik van Rijn began a third major chapter of theoretical development by merging the SBF model with an oscillatory working memory model and using concepts of oscillatory multiplexing to draw out the relationship between interval timing and working memory (Gu, van Rijn, & Meck, 2015). Later, the relevant oscillatory patterns during timing were shown using electrophysiological recordings from rodent striatum and cortex (Gu, Kukreja, & Meck, 2018). In addition, the SBF model was further extended by incorporating the SBF model into the ACT-R cognitive architecture (van Rijn, Gu, & Meck, 2014).

Warren and Bin Yin conducted lesion studies of the hippocampus in mice, but with more precise tissue damage than was possible in the rat work Warren had conducted in the 1980s, moreover they also included behavioural analysis of genetically modified mice (Yin & Meck, 2014). With Jessica Lake, Warren revisited the effect of dopaminergic drugs on timing, but in humans (Lake & Meck, 2013), and also examined the relationship between emotion and timing in humans (e.g., Lake, LaBar, & Meck, 2016).

Warren and Zhuanghua Shi used a Bayesian framework to revisit classic models of interval timing, revealing a new perspective for explaining the scalar property of timing and distortions in temporal memory (Shi, Church, & Meck 2013). This Bayesian approach could also explain previous findings of time distortions in Parkinson’s patients and aged individuals (Gu, Jurkowski, Shi, & Meck, 2016). Moreover, several review papers with Melissa Allman addressed temporal distortions in pathophysiological conditions (e.g., Allman, & Meck 2012).

More recently, Warren, Nicholas Lusk, and Elijah Petter examined the relationship between timing durations in sub-second and supra-second ranges, both with respect to the putative different roles of striatal and cerebellar circuitry (Petter, Lusk, Hesslow, & Meck, 2016) and in terms of how to best approach fitting data obtained from the same behavioral task (e.g., duration bisection), but different time ranges (Lusk, Petter, & Meck, 2020). This research complements earlier work with Sara Cordes (Cordes & Meck, 2014). Projects in progress with Elijah and Aryana Yousefzadeh extend this line of inquiry to a third brain area by infusing the striatum and hippocampus with muscimol, as well as asking more specific questions about the role of the cortico-striatal direct and indirect pathways in timed performance.

Warren’s continued interest in how temporal memories are stored in the brain led to a deep dive into molecular and biochemical mechanisms. In a highly promising application of a relatively new brain manipulation technique, Warren, his students and collaborators developed a paradigm that combines a novel licking task with optogenetics and quantitative behavioral analysis to permit dissection of how the basal ganglia support interval timing (Toda, Lusk, et al., 2017). A recent review article examined the role of synaptic plasticity and microtubule proteins as a potentially unifying mechanism for the acquisition and maintenance of temporal memories across varied timescales (Yousefzadeh, Hesslow, Shumyatsky, & Meck, 2019). At the time of his death, Warren was working on an invited review manuscript for Nature Reviews Neuroscience with May-Britt and Edvard Moser to provide a comprehensive understanding of episodic timing and interval timing and a more unified model of timing and time perception.

These review articles and search for unifying mechanisms of temporal processing constitute a critical aspect of Warren’s contribution to the field in the 2001-2020 era: Theoretical and review articles that sought to increase the audience for interval timing research, by making major findings more accessible, and by demonstrating the connections between interval timing and other domains of cognition and perception. In aggregate, these papers have been cited thousands of times, suggesting this strategy was successful. For many of these review papers, Warren worked with a co-author with whom he had not previously published. In some instances, these co-authors were already working directly on interval timing, but in other instances Warren sought out co-authors who were working on topics that he thought could be meaningfully connected to interval timing. In a few cases, Warren paired with a collaborator on multiple review papers (e.g., Melissa Allman, William Matthews, Hedderik van Rijn).

The topics reviewed from an interval timing perspective reflect Warren’s broad research tastes and belief that “timing is everything”. They included genomics [1], choline supplementation [2], neural circuitry [3], reaction time [4], neuropsychology [5], emotion [6], memory representation [7], neuroimaging [8], drug abuse [9], cognitive aging [10], neurochemistry [11], molecular mechanisms [12], Vierordt’s law [13], EEG/ERP/fMRI signatures [14], pathophysiology [15], Bayesian optimization [16], hippocampus [17], coding efficiency [18], psychopathology [19], clock properties [20], individual differences and ecological validity [21], oscillations and working memory [22], perception, attention, and memory [23], consciousness [24], social processing [25], neurobehavioral genetics [26], circadian rhythms [27], reinforcement learning [28], cerebellum [29], and signal duration [30].

Warren also increased the impact of, and audience for, timing research by organizing edited volumes. His first effort was a book, published by the CRC Press in 2003, entitled Functional and neural mechanisms of interval timing. He followed this up by guest editing/co-editing special issues of Cognitive Brain Research (2004), Brain and Cognition (2005), Frontiers in Integrative Neuroscience (2012), and Current Opinion in Behavioral Sciences (2016). These volumes share the feature of bringing together authors who investigate timing from a variety of theoretical perspectives and empirical approaches. It is also notable that while the book had 21 chapters, the Current Opinion in Behavioral Sciences special issue had 42 articles, and the Frontiers in Integrative Neuroscience special issue had 59 articles. Clearly, the field grew substantially during Warren’s career and he played an integral role in that growth.

Warren also was instrumental in the creation of the first formal timing community when he participated in the COST-ESF Networking grant “Time In MentaL ActivitY: Theoretical, behavioral, bioimaging, and clinical perspectives” (TIMELY), which was initiated and coordinated by Argiro Vatakis. He joined in most of TIMELY’s activities, giving talks at TIMELY meetings in Athens, Seville, and Tuebingen. Importantly, he served as an instructor at two TIMELY training schools: Groningen in 2011 and Corfu in 2013.

When the TIMELY networking grant ended, there was an obvious need to continue formal gatherings of the timing community. Consequently, Argiro and Sundeep Teki began working to establish a new academic organization for timing researchers. Warren threw his full support behind their effort and the Timing Research Forum (TRF), with Argiro and Sundeep as founding directors, came into formal existence in 2016. Warren was the keynote speaker at the first TRF conference in Strasbourg in 2017, where he presented a synthesis of his nearly four decades of research on the functional and neural mechanisms of interval timing.

In 2013, Warren, Argiro, and Hedderik van Rijn founded the first journal fully dedicated to timing research, and served as co-Editors-in-Chief. The initial thought was to name the journal Time Perception, but Warren immediately suggested Timing & Time Perception, referring to the 1984 edited volume by Gibbon and Allan. Warren pushed to make Timing & Time Perception an open access journal. From a two-issue inaugural volume, the journal has expanded to four issues per year, and has become, as its name implies and Warren desired, “a home for everyone in the timing family”.

Warren’s broad, integrative approach reflected his devotion to expanding the audience for timing research while bringing together its different branches. Indeed, his guidance and mentorship led to novel integrations of timing research with other domains such as decision-making, learning, memory, and computational modelling. He always made an effort to be supportive of anyone trying to enter the timing field regardless of their methods or the populations they focused on. He highlighted the value of their ideas even when they were not in full agreement with his own. Of course, he was confident that once they were part of the field he could convert them to his way of thinking.

Warren was an inspirational mentor, whether official or unofficial, to many scientists in the field of interval timing across multiple generations. The timing community is deeply indebted to him for his many and varied contributions. He will be missed.

Review Papers 2001 – 2020
(Meck, 2001) [1], (Meck & Williams, 2003) [2], (Meck & Benson, 2002; Lustig, Matell, & Meck, 2003; Meck & N’Diaye, 2005; Buhusi & Meck, 2005; Merchant, Harrington, & Meck, 2014; Van Rijn, Gu, Meck, 2014; Lusk, Petter, MacDonald, & Meck, 2016; Petter, Lusk, Hesslow, & Meck, 2016) [3], (MacDonald & Meck, 2004) [4], (Meck, 2005) [5], (Droit-Volet & Meck, 2007; Agostino, Peryer, & Meck, 2008; Cheng, Tipples, Narayanan, & Meck, 2016) [6], (Cordes, Williams, & Meck, 2007) [7], (Meck, Penney, & Pouthas, 2008) [8], (Williamson, Cheng, Etchegaray, & Meck) [9], (Balci, Meck, Moore, & Brunner, 2009; Lustig & Meck, 2011) [10], (Coull, Cheng, & Meck, 2011) [11], (Agostino, Golombek, & Meck, 2011; Yousefzadeh, Hesslow, Shumyatsky, & Meck, 2019) [12], (Gu & Meck, 2011) [13], (van Rijn, Kononowicz, Meck, Ng, & Penney, 2011; Kononowicz, van Rijn, & Meck, 2018) [14], (Allman & Meck, 2012; Allman, Pelphrey, & Meck, 2012; Gu, Jurkowski, Lake, Malapani, & Meck, 2015) [15], (Shi, Church, & Meck, 2013; Turgeon, Lustig, & Meck, 2016) [16], (Meck, Church, & Matell, 2013; MacDonald, Fortin, Sakata, & Meck, 2014) [17], (Matthews, Terhune, van Rijn, Eagleman, Sommer, & Meck, 2014) [18], (Allman, Yin, & Meck, 2014) [19], (Allman, Teki, Griffiths, & Meck, 2014) [20], (Matthews & Meck, 2014) [21], (Gu, van Rijn, & Meck, 2015) [22], (Matthews, & Meck, 2016; Teki, Gu, & Meck, 2017) [23], (Yin, Terhune, Smythies, & Meck, 2016) [24], (Schirmer, Meck, & Penney, 2016) [25]; (Yin, Lusk, & Meck, 2017) [26], (Agostino, Acosta, & Meck, 2017) [27], (Petter, Gershman, & Meck, 2018) [28], (Bareš et al., 2019) [29], (Lusk, Petter, & Meck, 2020) [30].

Warren’s complete list of publications may be viewed at:
https://scholar.google.com/citations?user=OFydajcAAAAJ&hl

Memories

A collection of pictures and videos of Warren can be found here. Please feel free to upload.

John Wearden, Emeritus Professor of Psychology at Keele University, UK:

I only met Warren maybe 5 or 6 times, and he was always very kind to me when we did meet. His death is a massive loss to the field, and a personal tragedy for Tina, his other family and close friends, of course. My only Warren Meck anecdote is a rather dull one, alas. We were both at a conference, in France I think. I had an old laptop, heavy, underpowered, squarish like a dark flattened brick. Warren arrived with a gleaming white Mac, slim yet with elegant curves, looking like a robot mollusc beamed from 20 years in the future. Unfortunately, the conference projector thought Warren’s laptop came from the future too, and resolutely refused to connect to it. Our hero wasn’t fazed. “I’ll just burn my talk onto a CD”. We all gasped. “You can do that?” Our own machines could barely read a CD, let alone write one. The deed was quickly done, and Warren’s talk was uploaded onto my machine, which was ancient enough for the projector to recognise it. I was happy to have been of service.

John C. Neill, Associate Professor of Psychology at Long Island University USA:

My deep and sincere condolences to Tina Williams. Warren and I became friends when we were undergraduates in the animal labs at UCSD, around 1976. As an undergraduate, Warren was uniquely self-reliant. Warren slept and cooked in an old VW bus, which he hid in the wild canyons near campus, and the campus security guys would sometimes roust him and force him to relocate. Warren generously helped me run my senior thesis pigeon experiments and we shared some text books to save money. Later, when we were graduate students, I was at Boston U and he was at Brown, we had time to talk about our new directions. He was continuing to study biological sciences as well as psychology, which helped his wide ranging studies in timing. I had no trouble believing him when he said that he routinely stayed up working all night. Later, I was fortunate to hear Warren and his wife, Tina and their student, give a talk about choline and learning at the Eastern Psychological Association. Multiple conversations about choline with them helped me to begin a new successful choline research project at Children’s Hospital, Boston. They introduced me to their choline colleague at Boston University, Jan Krzysztof Blusztajn, without whose collaboration we could not have been successful. This introduction led to a lot of new work showing beneficial effects of dietary choline supplementation in preventing or ameliorating seizure-induced brain damage. I also fed my wife lots of choline rich food when she was pregnant, and our twins turned out to be so smart that we have to give some credit to the choline hypothesis. In our most recent email correspondence, Warren was very helpful and encouraging, as usual, when I described some of my ideas about timing and music research, and I’ve received a small grant to work on these ideas. Warren and I only saw each other once in a while at conventions, but it was always good to see him and discuss research ideas. I liked to kid him about how much mileage he got with a simple timing procedure. I recently had the good fortune to talk with Warren’s graduate student, S. Aryana Yousefzadeh, at their poster which was presented at the 2019 Society for Neuroscience meeting, and I think their new and exciting work on microtubules is very important, not only for understanding timing, but for our understanding of learning, in general. See: https://www.frontiersin.org/articles/10.3389/fnmol.2019.00321/full

Patricia Agostino and Diego Golombek, Chronobiology Laboratory, National University of Quilmes, Argentina:

Warren was an excellent scientist and a truly exceptional person. When Patricia stayed for a few months as a postdoc in his lab in 2008, not only he mentored her in the area of interval timing, but also taught her about life. We remember him always cheerful, making jokes. Warren was absolutely essential for our lab in Argentina: he taught us about new temporal domains, and it is fair to admit that we were completely circadian-centered. He helped us start a completely new research, dedicated to temporal processing in the seconds to minutes range and motivation. Since then, we always kept in touch. He visited us in Buenos Aires in 2014. Warren was always present despite the distance, and when Patricia’s baby girl was born last year, he claimed he would certainly know her soon… He always cared about our family and loved ones. We will miss him deeply.

2008, Durham, NC 
2014, University of Quilmes 
2014, Buenos Aires, Argentina 

Daya S. Gupta, Camden County College, USA:

I remember Prof. Warren Meck from SFN meeting in San Diego in 2016 where I was fortunate to share the aisle with him.  I remember him as a kind and affable person. When I was being introduced to him, he promptly acknowledged my presence even though the introduction was not complete. I also remember how he was carrying food and drinks in an ordinary plastic bag for his students. I believe that these were some of his qualities that saw timing research grow under his guidance.

Rich Ivry, Professor of Psychology and Neuroscience, University of California, Berkeley, USA:

In the winter of 1981, I was an undergraduate at Brown University, doing a senior thesis in psychology on perceptual development.  The research project, as well as new friendships with members of my lab, led me to start attending Department colloquia and social events, and just spend a lot of time around the Psych building.  Early on that year, I remember taking note of this guy who was ALWAYS there.  Morning, evening, nighttime—it just seemed that every time I turned a corner, there he was, usually moving at a very fast pace.  Never did get the courage to say hello, but I did ask my lab mates about this whirling dervish, and learned that he was a post-doc with Russ Church, doing some “weird experiments about how rats tell time”.  Wasn’t until four or five years later when I was starting my own grad school timing research that I came to know the fruits of Warren’s labors: A chapter that appeared in Gibbon and Allan’s 1984 Timing and Time Perception volume.  My xeroxed copy of that classic was the first of many many Meck articles that were to form the thickest folder in my steelcase filing drawers.  Such elegant experiments, setting the standard for a true cognitive neuroscience of timing that advanced both our understanding of fundamental psychological operations and the underlying neural mechanisms.  A marvel to see that creativity and insight sparkle for the next 35 years.  And, over the years as I got to know him through conference interactions, come to appreciate that Warren actually could slow down, at least in body if not in mind, taking me aside to provide an insightful critique of a poster or talk from the lab, delivered in his mischievous, ironic style.  Grateful for having been a witness for so many years to this true scientific pioneer. 

Bon-mi Gu, Postdoctoral scholar, Berke Lab, Department of Neurology, UCSF, USA:

Warren was a great advisor to me. When I first joined his lab, he gave me a USB drive full of timing papers, and that was the beginning of the brainwashing (he often liked to use this word) of my understanding in timing and time perception. The brainwashing happened gradually and smoothly throughout my grad school time with him. His interest in timing and time perception was very deep and broad so I was able to have a fun and memorable journey of timing research under his guidance. Throughout the journey, he was a great model of a lifelong timing researcher with strong passion. He was so passionate that he often lost track of time when talking about timing. I remember one occasion when he made a joke at the end of a prolonged lab meeting saying that losing track of time is a character of many timing researchers.

In addition to the guidance in timing research, he always wanted to help and support the careers of his grad students. This gave me a very supportive feeling that I can reach him and ask advice whenever I needed help. I did not realize it until he was gone that I had a mentor who I was implicitly relying on a lot. From my understanding, his caring attitude made him pretty busy because he would take full responsibility when someone needed help. I remember when his father was sick, he traveled almost every weekend from NC to New York (or somewhere far like that) to take care of his father. During this time, he still cared in full about my research and other concerns. I think that his caring attitude was also expressed in his very precise and detailed memory about other people and events. I was often amazed by his detailed memory, sometimes he even reminded me of events that I told him about and had forgotten.

His passing is such a great loss which came too fast. I still have a bottle of Whiskey he gave me when I was graduating. I meant to open it when I become a fully independent researcher and it is so sad that he has passed before I open this bottle. However, I believe his spirit and his research will continue to live on through the people who remember him. May he rest in peace.

Argiro Vatakis, Assistant Professor, Department of Psychology, Panteion University of Social and Political Sciences, Greece:

Warren was a constant inspiration to me, through his work and his passion about timing.

Warren was instrumental in the creation of the first timing community as he encouraged me to apply for a COST-ESF Networking grant for the establishment of such a community. I was fresh out of PhD and feeling the need for a timing community but also feeling too timid to attempt such a step. So, I emailed Warren asking his opinion on the formation of such a community. I was not expecting a reply as my email was without a specific plan and he did not know me. He replied immediately, encouraging me to take the step, and offering his help if I needed it. His reply was enough to get me started. The grant was funded from 2010 to 2014 entitled “Time In MentaL ActivitY: Theoretical, behavioral, bioimaging, and clinical perspectives” but everyone used the acronym TIMELY.

Warren was present in most of TIMELY’s activities supporting the newly founded community. He was there in the 1st TIMELY Workshop held in Athens in 2010 with a talk on “Neural Features of Encoding, Maintenance, and Decision Processes in an Ordinality Task Requiring Temporal Comparisons” (enjoy his talk here https://drive.google.com/open?id=1wOKwwNA55XYvvQNz1YW-wS-U_UL4v_2F).This is when we finally met in person!

In 2011, he visited Europe twice for TIMELY, once for the very successful training school in Groningen on the “Psychophysical, Computational, and Neuroscience Models of Time Perception”, where he trained along with Trevor Penney a number of young researchers on EEG and research on time perception and talked about the Neurobiology of time (enjoy his talk here https://drive.google.com/open?id=19wkIRWmGwqdQne3lbxoCCIj0nMlKbWq1). His second visit was for the TIMELY EBBS Satellite Meeting in Seville on the “Neurobiology of Time Perception: from normality to dysfunction”, where he talked about the “Neurobiological Models of Interval Timing”.

In 2012, he joined TIMELY’s Workshop on “Temporal Processing Within and Across Senses” in Tuebingen, where he talked about the “New perspectives on Vierordt’s Law: Memory-mixing and sensitivity to feedback in temporal comparison procedures”. Finally, in 2013 he joined another yet successful TIMELY training school in Corfu on “Timing and Time Perception: Procedures, Measures, & Applications”, where along with Trevor Penney he gave a great theoretical and hands on session on “Clock, memory, and decision-making processes in duration-bisection and ordinal-comparison timing procedures”.

He also supported TIMELY by participating in all the publishing activities of the action with chapter contributions in Multidisciplinary Aspects of Time and Time Perception (https://www.springer.com/gp/book/9783642214776#otherversion=9783642214783), Time Distortions in Mind: Temporal Processing in Clinical Populations (https://brill.com/view/title/21751), and Timing and Time Perception: Procedures, Measures, & Applications (https://brill.com/view/title/26665?rskey=QENNZV&result=1).

Warren was also instrumental in the establishment of the first timing journal. The initial thought was to name the journal Time Perception but Warren immediately suggested Timing & Time Perception to be the most appropriate title of the first timing journal. The cover of Timing & Time Perception was also Warren’s idea, which was a painting by the artist Francois Pouthas that was given as a present to his mother Viviane Pouthas, who devoted most of her life working on timing and time perception. Warren envisioned Timing & Time Perception to be open source, he also envisioned a sister journal for hosting review articles only. This vision came true in 2014, however, busy schedules and limited funding and support did not allow for this open source journal to grow as we had envisioned. So it is a project to be continued.

The need to continue the timing community after TIMELY was completed was discussed many times until the establishment of the Timing Research forum (TRF) in 2016. Warren was the “initiator” for this new community given that one day he emailed me a paper by Sundeep Teki (https://arxiv.org/pdf/1512.00058.pdf), where Sundeep was pointing out the need for a timing community. Warren knew I needed someone to help with establishing yet another timing community and he had found the most appropriate person to pair up with me. He did not say it explicitly but he knew that if I read the paper I would act on it.

Warren was again there in every step offering his support, ideas and materials related to timing. TRF’s collection of book and special issue covers was curated by Warren (see here http://timingforum.org/publications/books/ and here http://timingforum.org/publications/special-issues/). And of course, he was there in all TRF conferences. In TRF1 in 2017 in Strasbourg he gave us an excellent keynote talk on the “Functional and Neural Mechanisms of Interval Timing”, while in TRF2 in 2019 in Queretaro he organized a great Symposium on the “Differential Roles of Cortico-Basal Ganglia and Entorhinal Cortex-Hippocampal Circuits in Timing and Episodic Memory”.

He will be very much missed. He will continue to inspire us through the enormous wealth of knowledge he left behind for all of us.

Fuat Balcı, Professor of Psychology, Timing & Decision Making Lab, Koç University, Istanbul, Turkey:

Warren will be deeply missed by many generations of scientists.

I first met Warren at Columbia University/NYSPI in 2003; at the time, I was a graduate student at Rutgers University and I was visiting the Temporal Cognition Unit on a weekly basis (Warren was a visiting scholar at Columbia University/NYSPI). I learned a lot from my interactions with Warren. He made very inspiring and motivating suggestions for my research; these conversations have had a long-lasting impact on my thinking and work. We kept in touch at conferences and always had many things to talk about regarding our ongoing studies and recent developments in timing research. We co-authored our chapter on “Timing Deficits in Aging and Neuropathology” during this period; it was a great pleasure working with Warren on this chapter. We then met at the TIMELY Training School on Psychophysical, Computational and Neuroscience Models of Time Perception. This was a great gathering during which I had many opportunities to talk to Warren and get his feedback on recent developments in my research agenda; he was always very attentive and supportive. During this visit, Warren, Trevor Penney and I rented a car and made a small excursion to the dikes and explored the windmills. Some of the following photos were taken during this excursion. As we were driving back to Groningen, we saw a giant pink dove figure in the backyard of a property. We “had to” take a closer look at it and we did, only to find out that there was another surprise in that backyard, a retired fighter jet (F-104G).

The Netherlands (dikes), 2011
The Netherlands (dikes), 2011 (Photo contributed by Trevor Penney)
The Netherlands (dikes), 2011
The Netherlands (inside a windmill), 2011
The Netherlands (Giant pink dove figure and F-104G Starfighter), 2011
The Netherlands, Groningen, 2011

I hosted Warren and Tina Williams at Koç Univesity, Istanbul in 2012. Their visit was very impactful and helped me bring together many researchers, who are interested in timing. This ultimately helped build a sense of community for timing researchers in the country. During this visit, we had long discussions with Warren about the Timing & Time Perception journal as we were watching the Bosphorus from a location a few kilometers south of the presumed location of Symplegades. We discussed how the journal can consolidate the efforts of timing researchers and contribute to the formation of a timing community. I can still remember how enthusiastic and excited Warren was about the journal. His enthusiasm was truly contagious as it has always been.

Istanbul, Turkey 2012

Shortly after, we came back together in Corfu (Greece) for the TIMELY School on “Timing and Time Perception: Procedures, Measures, & Applications” that Argiro Vatakis, Georgios Papadelis and I organized. During the entire duration of the training school, Warren paid very close attention to the questions of junior researchers and graduate students and very generously provided guidance to them. The picture below was taken during Warren’s interaction with graduate students during the TIMELY School at Corfu; Yes, he chose to sit in the middle of the room.

Corfu, Greece 2014
Corfu, Greece 2014
Corfu, Greece 2014

Warren, Tina, Trevor, and I made another excursion during this trip. At times the back roads on the island were blocked by free-range goats. Warren was the only one to get out of the car to herd the goats away from the road. We even drove up to the Mount Pantokrator to have a nice view of the island. Well, we could hardly see each other because of a very heavy fog but we could certainly hear each others’ laughter. I can never forget our nice outings and suppers in Corfu that were spiced with friendship and science.

Our last interactions were in Mexico, Queretaro during TRF2. During a bus ride from the campus to the hotel, Warren introduced a new timing researcher to me. He was highlighting the ways in which our research could support each other. This was a very familiar scene for me; Warren has always been very supportive of researchers and helped integrate them into the timing community. Whenever I saw Warren interact with graduate students and junior researchers, I could always see the same enthusiasm on his face that I witnessed back in 2003 while interacting with him as a graduate student. We will miss you a lot, Warren.

Melissa J. Allman, Assistant Professor of Psychology, Michigan State University, USA:

I was shocked and deeply saddened to learn of Warren’s death, what a devastating and enduring loss to the timing and time perception field at large, and to his many colleagues and collaborators, former and current students, and of course, Tina. My first contact with Warren was memorable for me, even though it was via email—I had written a K99/R00 proposal that I sent him on timing in autism as I wanted his permission to use a figure from one of his publications in my grant (I thought it might look more impressive to reviewers if it said ‘with permission’ in the legend, given his timing legend status). To my absolute horror, when checking my sent box to make sure it had gone through, I realized my over-zealous spell-checker had corrected his name to Neck, throughout. When he replied (with what I came to find out was his usual) kindness, wit, and generosity, not only did he grant me permission to reproduce his figure, he offered to be a co-mentor on my grant, and for me to have weeks-long study visits at Duke with him and his lab (I had landed a legend on my grant!). From that first exchange of emails, we ended up having many successful writing collaborations over ten years; he had a huge impact on my career. Warren was a good mentor as he was supportive, could use you in ways that benefit you both, was a rigorous scientist and was inspiring. Much like he was for me, perhaps his legacy is as ‘Father Time’; having an enduring presence, steering the field, and providing a solid foundational base for much of what we know about timing and time perception at the behavioral and neuropharmacological levels, upon which new lines of investigation grow. He and his work will be missed and remembered.