Hormones and Behavior 66 (2014) 743–758

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In memoriam

Jay S. Rosenblatt (November 18, 1923–February 15, 2014)

Jay S. Rosenblatt was a pioneer in the study of the neural and endocrine bases of parental behavior in mammals. Jay's research on the mechanisms underlying maternal behavior in cats, rats, and rabbits, and the development of behavior as a synchronous interaction between caregiver and offspring through weaning, created a genre of international maternal behavior research. He created unique experimental preparations and paradigms, many of which continue today as standards in the field. In addition, in complete opposition to the prevailing view in the field at the time, he demonstrated that young animals were fully capable of learning those things vital for their survival. These studies resulted in over 160 research publications and many review chapters. Raised in the Bronx, Jay never intended to become a researcher. His first interest was painting and he began taking lessons as a teenager before starting his undergraduate studies at the College of the City of New York. After serving in the Army, he enrolled at New York University to complete his studies in animal behavior. Six years after earning his Ph.D., Jay went on to train as a psychoanalyst at the Postgraduate Center for Mental Health in New York and established a private practice, which he maintained for 25 years while

http://dx.doi.org/10.1016/j.yhbeh.2014.09.006

also working as a researcher at Rutgers and continuing his painting. Jay received significant awards in recognition of his research contributions: the first Daniel S. Lehrman Lifetime Achievement Award from the Society of Behavioral Neuroendocrinology in 2006, the Senior Investigator Award from the International Society for Developmental Psychobiology in 2007, the Honoris Causa Doctorate from the University of Gotebörg Sweden in 1987, and the Doctor Honoris Causa from the National University of Distance Education in Madrid, Spain in 1997. The scholarly journal, Developmental Psychobiology, paid him an unusual honor: a special issue in January 2007 in recognition of his contributions to the study of animal behavior. He was a longtime co-editor of the scholarly series, Advances in the Study of Behavior, and a fellow of the American Association for the Advancement of Science. Jay had an incredible 40 plus consecutive years of federal funding to support his research. Jay began his career at Rutgers in 1959 and served as Director of the Institute of Animal Behavior on the Newark Campus from 1972 until 1989 and as Acting Dean of the Graduate School in the late 80s. In 1984, the alumni of the Institute held a Fig. 2 and Fig. 3 pertain to this conference and published a special volume by the New York Academy of Sciences in Jay's honor. Jay retired from Rutgers University in 2005 as the Daniel S. Lehrman Professor of Psychobiology. He continued his scholarly activities until shortly before his death this year at the age of 90. Last year, at the age of 89, he had largely completed a 246 page manuscript entitled, “The Evolution of Parental Behavior among the Vertebrates: A Review.” Jay was a wonderful, warm, sensitive, caring, mentor and role model to young investigators. He had a profound impact on launching the academic research careers of many young women and men. Harold I. Siegel JSR — remembrances In 1969, I was a fourth-year undergraduate at Rutgers University (New Brunswick Campus) working on a study of the dietary selfselection patterns of pregnant and lactating rats with Alan Leshner, then an advanced graduate student in George Collier's lab. As pregnancy progressed and then during lactation, the rat mothers chose to eat more from the food holder that contained protein (and other ingredients) but no carbohydrates instead of from the holder that contained carbohydrates (and other ingredients) but no protein. This was my first such laboratory experience and I was thrilled. However, what I found even more interesting was watching the mothers take care of their pups; the near constant attention to the young that included nest building, licking of the pups, crouching over and retrieving them when necessary. At that point, I knew that I preferred a research career over a clinical one,

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and I knew that I wanted to pursue the study maternal behavior. My knowledgeable and trustworthy sources recommended in no uncertain terms that I apply to the Institute of Animal Behavior (IAB) and sign on with Jay Rosenblatt. I sent in my application and waited. I heard nothing. I began calling twice per day asking to speak to Dr. Daniel Lehrman, Director of the Institute. The secretary was impenetrable. Finally, Dr. Lehrman unexpectedly answered the phone and rather abruptly told me to show up one day the next week for an interview. On April 1 (April Fool's Day), 1969, I parked my car in a commercial lot in Newark, NJ. Unfortunately, no one told me that the Institute had moved across town. I made my way to the new Institute and eventually (no one told me to take the elevator instead of trying to enter the IAB through the locked stairwell doors) was led to a room where I met the other interviewees: John Buntin, David Crews, Joan Herman, and Rae Silver. Dr. Lehrman arrived with four folders; mine was missing (and as far as I know, never found). Nevertheless, I was permitted to participate in the interview process and met individually with faculty as I was directed. It was mid-afternoon and I had some free time and decided to make certain that I met with Jay Rosenblatt, the primary reason for applying. No one told me that he left most days in the middle of the afternoon. It was curious to me that a faculty member would just leave, and I then felt really dubious about any possibility of admission. I learned in the fall of that year, after I began my graduate studies at the Institute along with John, David, Joan, and Rae, that Jay left Newark mid-afternoon to return to his New York City apartment to see his psychotherapy clients. How I managed to be accepted, I do not know; how my life was so positively affected, I do know. My first year with Jay was not really with Jay. He had a number of more advanced and very bright students including Joseph Terkel, Alison Fleming, Bob Woll, Neil Whitworth, and Natalie Freeman. Actually, I was quite lucky and learned a great deal from each. Perhaps most valuably, one day I asked Alison how one knew when one had done enough for a dissertation. She told me very clearly that all I had to do was to tell Jay that I had done enough. A couple of years later I discovered she was correct. The atmosphere at the Institute was ideal for the discussion of ideas. Many would gather early each day around a long table and we would talk about our own studies, those of others, and those we read about. Energized, we left to our respective offices and labs. Jay and I would speak most mornings; I not only felt nourished by his words but also I felt as if we could conduct the best possible studies. Our studies did go well, as did those of the others in Jay's lab. Jay had developed his pregnancy termination model that permitted experimental control over late pregnancy events. Rats in the latter stages of their pregnancies were either hysterectomized (H) or they were hysterectomized and ovariectomized (HO). The H animals when presented with foster pups 48 h later showed short-latency maternal care similar to what occurs after natural termination. The HO animals eventually behaved maternally but it took them significantly longer. The question was thus, what were the ovaries doing following the H procedure alone that contributed to the onset of care? In those days, other researchers had recently turned their radioimmunoassay procedures to measuring ovarian steroids during pregnancy. The picture at the end of pregnancy showed that estradiol was increasing and progesterone was decreasing. It made sense to incorporate this information and we began to inject estradiol in increasing doses and progesterone in decreasing doses beginning immediately after the HO surgery. The doses chosen were initially quite arbitrary and after a variety of hormone treatments, we were able to demonstrate that estradiol alone resulted in short-latency maternal care; progesterone treatment during the immediate post-operative period could actually slow the onset of the behavior. Jay had our test cages custom made. They were not the usual size rat cages and instead they were much larger. Larger cages used up so much shelf space that sometimes they limited the number of subjects we could run simultaneously. One day, we were in a hurry to empty the

subjects from several cages in order to provide the space for new ones. In haste, a number of the H animals who had already become maternal were placed into the same observation cage. In response to the stimulation from the other rats (all females), some of them began to show lordosis. Their pregnancies had been terminated, their ovaries were secreting estrogen, and once actively maternal, progesterone levels increased and the animals entered “normal” post-partum estrus. It was an accidental observation that again proved the value of Jay's pregnancy termination model and paved the way for further understanding of the hormonal onset of the behavior and led to many of our subsequent studies and those of others that included the initial investigations into the role of hypothalamic steroid receptors. During my graduate years, Danny Lehrman passed away (1972) and Jay became the Institute's second ever Director. I completed my degree and after a post-doctoral stint, I returned to the Institute to continue my studies on rats and hamsters with Jay and others including Bob Bridges, Henry Szechtman, and Peggy McCarthy. For various academic and political reasons, the Institute was merged with the Department of Psychology, making for an initial uneasy transition that is now water under the bridge. From this point forward, Jay and I were members of Psychology along with most other IABers and the cognitive and clinical psychologists. It was also at this time that my father passed away and in many respects Jay intentionally or not served as my surrogate father. Except for some time out for a triple bypass, Jay continued to be the academic he had always been. He eagerly sought out discussions with anyone willing to respond. He demonstrated a desire to learn about the other faculty's research. He often told me that when he visited other departments, he believed that he came to know more about the faculty's work than their own colleagues did. I believed him. He attended colloquia in Psychology and in the Center for Molecular and Behavioral Neuroscience. After changing my research to adult attachment including a major study of pedophiles, Jay continued to listen and offer compliments and suggestions. Jay officially retired in 2005 and continued to spend more time on campus than some of our full-time faculty. The weather might be terrible but if that was a day he was expected to be in his office, he was there. When I became Chair, my office was half-way between the entrance to our building and his office. He stopped to chat on each of his mornings. I knew that the stopover permitted him to rest and catch his breath before moving down the hall. He had told me on a number of occasions that Danny Lehrman did something similar, stopping for some non-health related reason that was actually to help him catch his breath. Jay never let on to Danny that he understood the temporary time-outs and neither did I. I looked forward to our early morning chats that covered an enormous range of topics including some psychological insights about some of those we both knew. He spoke about Gilda, his first wife, who passed away, and about Pat, his then current wife. He spoke with love and admiration. The same was true for Danny and Nina, his children and grandchildren. During our final several visits, Jay offered me two of his paintings, both dark and sad and representing the apocalypse (now on our dining room wall) and a family's learning about the death of another family member (now on my office wall). In both cases, the family members are huddled together providing support during such difficult times. Our topics of conversation during these last visits included Jay's recounting of some of his professional and personal regrets over his lifetime and his growing sensitivity to the plight of others in the world. I was taken aback. As naïve as it sounds, I would not have believed that this man I thought so highly of would have some of these questions and doubts. The mentor–student relationship can provide an absolutely wonderful bond. I had the very best. Thanks, Jay. Harold I. Siegel Jay Dedicated to Pat, Gilda, Danny, Nina, Olivia, and Suzanna

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Jay Rosenblatt has served many roles in my life. He was my PhD supervisor and scientific mentor. He was a teacher and a colleague. He was a friend and confidant and even sometimes, a therapist. And now that I am dabbling in the arts myself, I appreciate better, Jay, the artist and critic. As teacher and supervisor at the Institute of Animal Behavior, Jay spent many hundreds of hours in the seminar room, in his office, in the lab, while driving home to NYC — talking with us, his students, about his ideas and ours. He helped us design studies, kibbitzing along the way as we struggled with one confound or another; he critiqued our writing and helped ‘grow us’ into professionals. He supported us against external academic challenges and taught us how to gracefully accept criticism and even rejection. Jay's thinking and his gentle steady style and hand provided a role model for how to mentor. Jay was also the consummate scientist, and was passionate about the questions he asked and the answers he found. He would take any opportunity to talk science and to provide subtle, often not obvious explanations for psychologic phenomena. We, his students, spent many hundreds of hours at conferences, over the phone, by email talking science with Jay. Imagine, for over 45 years! And even in the week before he died when he was already quite weakened, we continued the conversation and spent a good 1/2 hour at his home talking about his scientific life, which studies he felt to be a contribution and which he wished he had followed up; what his Eureka moments were; and what studies just didn't work out. He knew that he had made substantial contributions to his science, but until the end he was modest about his accomplishments, too modest. He didn't quite accept that he had provided the most used framework for the analysis of maternal behavior over the past 50 years. In 2006 I wrote a chapter entitled “Three Faces of Jay S Rosenblatt: as Scientist, Painter, and Psychotherapist” for a Developmental Psychobiology volume that included papers and studies that grew out of, or interfaced with, Jay's framework and thinking. In preparation for that article I interviewed Jay extensively in Central Park in New York, I reread many of his scientific papers, especially his earlier ones, and I visited him and his family in their home. What follows is taken from that paper — albeit, considerably shortened. It provides some of the chronology and ontogeny of his scientific life, his goals, and his accomplishments. This was one of his ‘faces’. For a look-in to the others, the painter, the psychotherapist, and, of course, the father, grandfather, and husband, see Fleming (2006). If Jay objected to any interpretation of his life provided in this bio, he didn't mention it, and I take this to mean that it doesn't deviate too much from his truth. But let me start with the beginning of his scientific life. It didn't begin when he was a youngster, as with many biologically-oriented scientists. In fact, there were few indications from his earlier life that Jay would go into science. His favorite subjects in school were art and “the social sciences attracted (him) more than the sciences (which [he] feared a bit).” Between 1943 and 1945 Jay was in the army, stationed in various parts of Europe including several places in England, in Paris, and Liege, Belgium. In 1946 he entered NYU to complete his training. The years between 1945 and 1953 were pivotal to Jay's scientific career. At NYU he met T.C. Schneirla whose work and theoretical position were clearly reflected in Jay's own subsequent work and thinking. Schneirla convinced Jay that having a “firm grounding in animal behavior (would provide) the broadest basis for understanding human behavior in addition to being of intrinsic interest.” Jay describes a conversation between the two where Schneirla said “well you are not going to get rich studying animal behavior” to which Jay responded “well, I was prepared not to get rich becoming a painter, so I might just as well not get rich studying animal behavior.” And so Jay started his PhD work nominally with Schneirla but also with Lester Aronson studying the role of hormones and experience in the organization of sexual behavior in male cats (Rosenblatt & Aronson, 1958). It was at the American Museum of Natural History (AMNH) on a grant with Schneirla that Jay began his first foray into the study of mother–young interactions in cats. During that time he became very interested in early learning by the kittens and with Gerry Turkewitz

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who was a student, established that kittens develop home orientation by days 3–4; at the same time he discovered that kittens develop nipple position preferences on days 1–2 of life. This work was important because it was counter to the belief based on more traditional conditioning paradigms that animals cannot learn at such young ages; it showed that by using ‘natural’ species–characteristic situations, common now in the work of Jeff Alberts and others, experience and learning can be shown to occur very early in life (Rosenblatt, 1983). Earlier, as a teacher at NYU, later at City College as a TA, and then as Assistant Professor at CCNY, Jay became close with Herbert Birch, a human developmental psychologist, who for a period constituted another important influence in Jay's intellectual development. Birch worked on development in newborn babies, providing Jay with an outlet for his interest in human development. It was during these formative years in New York City that Jay met Danny S. Lehrman. This was a friendship that lasted until Danny's untimely death in 1972. Jay and Danny Lehrman were graduate students together at NYU and also were colleagues at CCNY as assistant professors, as well as associates of the Department of Animal Behavior at AMNH where both of them did their doctoral research. Danny was the more advanced than Jay in animal behavior as an expert ornithologist who had already published a research article at the age of 17, and was becoming world renown because of his critique of Konrad Lorenz in 1953 (he received his PhD in 1954!). Says Jay of this relationship: “I learned a lot from Danny but most of all we were warm friends who could talk to one another about the most important things in our lives.” This affection is reflected in Jay's biography of Danny written for the National Academy of Sciences (Rosenblatt, 1995; Silver & Rosenblatt, 1987). The 1950s was a difficult period in US history, during the McCarthy era and Jay and a number of his colleagues (Max Hertzman) were fingered as persons of interest by the House Un-American Activities Committee (HUAC), an ‘interest’ that was conveyed to the CCNY administration, which was followed by the nonrenewal of his contract. Jay soon thereafter moved to Rutgers University to join Lehrman in his newly created Institute of Animal Behavior in Newark, New Jersey. Jay began his work on rat maternal behavior after arriving at Rutgers. This work was what he spent most of his intellectual energy on for the next 50 years (Jay retired in 2005). It is not possible to discuss each and every one of the 150 + papers and chapters that Jay has written in his 60+ years as a scientist. I will, however, describe two of the primary themes that have grown out of his work and the primary influences he has had on the field and on his students. As Jay himself pointed out, “most theorists are known for one or two ideas that formed the framework of their thinking.” In Jay's case there are two and, maybe, three, principle ideas that have come to be identified with his perspective and have had an impact on the field in general and on the direction of my own work, in particular (for overview, see Corter & Fleming, 2002; Fleming & Li, 2002; Numan, Fleming, & Levy, 2006; Moore, 1995; Stern, 1990). Not all work done at the Institute of Animal Behavior or by his students and colleagues over the years are discussed below. For this I apologize. Instead, emphasis is given to the ‘earlier’ studies, and to the intellectual environment that existed when I was a student in the late 1960s and early 1970s and which were formative in my own thinking. 1. The first major contribution to the field was his recognition that given the right ecologic and naturalistic contexts one can demonstrate learning by the neonate at ages that are considerably younger than believed possible at the time this work was first done in 1960s (Rosenblatt, Turkewitz & Schneirla, 1969). The sensory modalities recruited for this early learning were initially single modalities, thermal, and tactile, but through associative processes, came to depend heavily on the olfactory sense and eventually became truly multimodal. Jay's early kitten work, illustrating these processes and described above, provided the groundwork for an entire field of study that focuses on learning within naturalistic contexts and by neonatal

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animals (Rosenblatt, 1971; Rosenblatt et al., 1969). This idea, tested well before its time, is reflected in more recent work called ‘the constraints on learning.’ Although Jay is most famous for his work on rat maternal behavior, which he started when he joined Danny Lehrman's new Institute of Animal Behavior in 1958, his first work on kittens (started in 1954 at the AMNH) really epitomizes his approach to the organization of behavior and the influence of Schneirla's Approach/Withdrawal theory (Schneirla, 1952; Schneirla & Rosenblatt, 1961), namely: that to understand development one must understand the transition between dependence on basic sensory–motor reflexes which occurs first, to the development of affectively based perceptual–motivational relations. Hence, through learning, simple responses to primary stimuli (thermal and tactile) that vary along the intensity dimension become approach or active withdrawal from affectively laden multimodal stimuli (often with the addition of olfactory information; Rosenblatt, 1971). When I asked Jay what was the most exciting personal moment in his career — a rather unfair question, I realize — he described it was these kitten discoveries, which were in fact, quite serendipitous. While weighing kittens on a daily basis he had noted and later set Gerry Turkewitz to study, that whenever kittens were replaced back into their home environments (after removal of the mother), the kittens would return to the home corner and do so more rapidly each day. Since the kittens seemed to know where to go, despite having no vision, this could only be based on early learning of the olfactory characteristics of the area where mother normally nurses her young (Rosenblatt, 1983; Rosenblatt et al., 1969). He says of this work and the kitten work that followed, “It seemed to me all the problems of early development could be studied in the development of home orientation of kittensdevelopment of sensory capacities and developmental transition in the use of sensory systems, development of motor capabilities, and the effect on sensory system use (when kittens rise off the floor (crawling to walking) they cannot use olfactory stimuli as well or as continuously and need to shift to vision with olfactory support, etc.), transition from the use of socially conditioned stimuli (nest odors) to social stimuli (mother), development of learning and cognitive structures (internalization of pathtaking) and of course, their emotional development indicated by their distressed vocalization and its termination” (Rosenblatt, 1983). 2. The second set of ideas or themes that have come to be identified with the Rosenblatt framework are reflected in the concepts of behavioral transitions in the maternal behavior cycle and behavioral synchrony between mother and young. Since these concepts have had such an impact on my own work and are by now part of the vocabulary of the study of maternal behavior (‘onset vs. maintenance’), they will now be discussed in some detail (Rosenblatt, 1970). Although the study of maternal behavior is not primarily about early development, Jay treated the phases in the maternity cycle much as he would any developmental problem, as a series of developmental transitions. He became very interested in the phenomenology and then the mechanisms that mediated the development of maternal responsiveness from mating through pregnancy, to pregnancy termination, through the postpartum period, and into and through weaning. Each of these phases was characterized, for each he established the role of sensory factors, the associated physiological changes, the feedback effects of behavior, the role of endocrine factors, and then the role of shifts in hedonic and affective mechanisms. For each phase the mother undergoes there occurs a synchronized set of behavioral and physiological changes in the offspring. While in his rodent work, the emphasis was on the mother, the developmental status, and needs of the offspring changed accordingly, so that mother and offspring were mutually adapted in their behavior to one another. According to this view maternal care begins before the young appear, with conception. Once mating and pregnancy have taken place, the mother to-be experiences endocrine changes that alter her behavior and psychology. She shows enhanced nesting

and self-grooming behaviors, she comes to restrict her movements and reduce activity level, focusing instead on a particular nest site; she changes her eating preferences and behavior, with increases in preference for needed nutrients; and she shows variations in emotional behavior, changes in perception, and comes to attend to some cues over others (Lott & Rosenblatt, 1969; Rosenblatt, 1980). The next transition in the maternity cycle that Jay and his students have studied most intensively occurs at the end of pregnancy and was first described in a chapter by Rosenblatt and Lehrman (1963). “The transition between the onset of maternal behavior and its maintenance is a powerful one; with onset being mediated by the hormonal changes occurring toward the end of pregnancy and at parturition.” While Jay did not know which hormones “turned on” maternal behavior initially, his 1972 studies with Joseph Terkel on the role of blood borne factors in the functional parabiotic manipulation showed that the relevant factors were present during the last 48 h of gestation (Terkel & Rosenblatt, 1968, 1972). These studies with Terkel were followed up by studies by other students and postdocs working at the Institute of Animal Behavior. These described the pregnancy effect in which mothers undergo elevations in maternal responsiveness across pregnancy, peaking close to parturition. By means of the famous (infamous) hysterectomy–ovariectomy (H–O + E) endocrine manipulation that came to be associated with the Rosenblatt lab, at various points throughout pregnancy, Jay and his students (especially Harold Siegel) established the importance of midpregnancy endocrinology and the subsequent decline in pregnancy progesterone and rise in estradiol for the onset of maternal behavior (Siegel & Rosenblatt, 1975b,c, 1980). These studies lead to a recognition that estrogen priming is also essential to the later discovered activational effects of oxytocin and prolactin in the initiation of maternal responsiveness. The third phase in the maternity cycle, “the maintenance phase,” has, until recently, received very little attention. However, again Jay did some landmark experiments on the role of experiences acquired during the postpartum on the subsequent expression of the behavior at a time when the parturitional hormones were no longer playing a role (Rosenblatt, 1980). These experiments suggested the existence of a sensitive period for the long term effectiveness of a postpartum experience with young; it also suggested a role for different sensory modalities in the experience. Work initially by Bridges (1975), Bridges et al. (1977), Cohen and Bridges (1981) and then by our and other laboratories (summarized in Fleming & Li, 2002; Numan et al., 2006; see also Stern, 1989, 1990, Stern, Yu, & Crockett, 2002) followed up on these early studies and we began to explore the role of the expression of the behavior per se and of somatosensory versus olfactory experiences with the pups in the maintenance of the motivation to mother. We illustrated the importance of the timing and duration of the maternal experience, and, as with other forms of learning, the importance of the interval during which pups are not present on responsiveness at test (the retention interval). These studies started by Jay show that reproductive behaviors like maternal behavior may be species-typical and relatively stereotyped in form, but they are nevertheless subject to the influences of experience and learning and exhibit considerable flexibility in when they are expressed and with what intensity. These behavioral flexibilities are mimicked by flexibility in brain function and structure. The end of the maternity cycle, that of weaning, was also of considerable importance to Jay but it remains to this day the least studied phase (Reisbick, Rosenblatt, & Mayer, 1975). Jay's early work on weaning by a mother cat of her kittens is a true classic and describes behavioral synchronies between mother and kittens, where the mother cat actively withdraws from her growing and proactive litter prior to weaning (Williams, Hall, and Rosenblatt. 1980), providing a facile analogy for our own experiences as parents of teenage children! In short, Jay and his colleagues and the IAB explored the psychobiology of maternal behavior and aspects of infant learning from many angles, at multiple analytic levels, using multiple technologies and techniques, as

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they became available, and from a proximal, functional, developmental, comparative and, more recently, evolutionary perspectives. He and his colleagues were among the first to study maternal behavior from the perspectives of its humoral and hormonal bases; he explored the role of sensory factors, starting with somatosensory stimulation of mammary gland development during pregnancy derived through self-licking and the role of chemosensory cues in its organization and regulation. His work on experiential factors (which harks back to his studies on the role of experience in the mating behavior of male rats following castration) is legion and begins with his famous chapter with Danny Lehrman in 1963 (Rosenblatt, J.S., & Lehrman, D.S., 1963). Jay's students and postdocs also explored the role of neural and neurochemical factors and led by Michael Numan, together with Barry Komisaruk, he was among the first to demonstrate the importance of the medial preoptic area (Numan, Rosenblatt, & Komisaruk, 1977). This work on neural control of maternal behavior, starting in 1975 was very fruitfully followed up over the years by other students and postdocs (Felton, Linton, Rosenblatt, & Morrell, 1998). Finally, in the late 1980s Jay, Harold and their students ventured into neuro-molecular work, establishing for the first time a role for nuclear estrogen receptors in the MPOA in the regulation of maternal behavior (Giordano et al., 1989, 1990, 1991). In fact, the analysis of the neural and neurochemical mediation of maternal behavior produced as much as 25% of all research papers coming out of the Institute of Animal Behavior/Neuroscience Institute at Rutgers. In Jay's more recent years he was working on a book on the evolution of parenting. And although it was a struggle, everyday he would work on it, but he couldn't quite decide what constitutes the phenotype that is of relevance to the evolution of parental behavior. Is it the behavior? The physiology? The overall social organization of the different orders and species? He struggled with these ideas until a few weeks before his death. However, at the age of 89 he had another eureka experience and here I quote from a recent (2013) email from Jay. “I had an insight about how to organize the material of my review and my thinking about the evolution. Instead of directly characterizing the PB as evolutionary sequences I now believe that the backbone of the evolution consists of the evolution of the reproductive physiology (sexual and parental) in the various vertebrate classes of which there are basically three patterns found in the fishes– amphibia, reptiles–birds, and monotremes–marsupials–eutherian mammals. The physiology provided the platform on the basis of which PB initially arose in the bony fishes as an aspect of mating behavior (egg-scatterers and related pattern). Further evolution of PB required elaboration of adaptations to specific features of the breeding habitat in each of the clades of fishes. The evolutionary sequences can be traced in detail in many fish species and this is the only true direct analysis of the evolution of PB. All other “evolutionary analyses' are derived but no less valid if recognized as such.” Although he was not able to finish this analysis, he invited anyone who was interested, to take on the task — a tall order!” So this describes Jay's history and thinking, especially during his earlier years at the IAB, from the perspective of a lifelong student. We miss Jay. He has had an enormous impact on our own careers and that of our scientific children, and theirs — and for that we are forever grateful. Alison S. Fleming References Bridges, R. S. (1975). Long-term effects of pregnancy and parturition upon maternal responsiveness in the rat. Physiology and Behavior, 14, 245–249. Bridges, R. S., Feder, H. H., & Rosenblatt, J. S. (1977). Induction of maternal behaviors in primigravid rats by ovariectomy, hysterectomy, or ovariectomy plus hysterectomy: effect of length of gestation. Hormones and Behavior, 9, 156–169.

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Cohen, J., & Bridges, R. S. (1981). Retention of maternal behavior in nulliparous and primiparous rats: effects of duration of previous maternal experience. Journal of Comparative and Physiological Psychology, 95, 450–459. Corter, C. M., & Fleming, A. S. (2002). Psychobiology of maternal behavior in human beings. Handbook of Parenting. In M. H. Bornstein (Ed.), Biology & ecology of parenting (Vol. 2, pp. 141–182). New Jersey: Lawrence Erlbaum Assoc. Fleming, A. S., & Li, M. (2002). Psychobiology of maternal behavior and its early determinants in nonhuman mammals. Handbook of Parenting. In M. H. Bornstein (Ed.), Biology & Developmental Psychobiology. http://dx.doi.org/10.1002/dev Fleming, A. S. (2007). The three faces of Jay S. Rosenblatt. Developmental Psychobiology, 49(1), 2–11. http://dx.doi.org/10.1002/dev.20193 Giordano AL, Siegel HI, Rosenblatt JS.(1991) Nuclear estrogen receptor binding in microdissected brain regions of female rats during pregnancy: implications for maternal and sexual behavior. Physiol Behav. 1991 Dec;50(6):1263–7. Giordano AL, Ahdieh HB, Mayer AD, Siegel HI, Rosenblatt JS.(1990) Cytosol and nuclear estrogen receptor rats after steroid priming: correlation with maternal behavior. Horm Behav. 1990 Jun;24(2):232–55. Giordano AL, Siegel HI, Rosenblatt JS. (1989) Nuclear estrogen receptor binding in the preoptic area and hypothalamus of pregnancyterminated rats: correlation with the onset of maternal behavior. Neuroendocrinology. 1989 Sep;50(3):248–58. Lott, D. F., & Rosenblatt, J. S. (1969). Development of maternal responsiveness during pregnancy in the rat. In B. F. Foss (Ed.), Determinants of Infant Behavior IV (pp. 61–67). London: Methuen. Moore, C. (1995). Maternal contributions to mammalian reproductive development and the divergence of males and females. In P. J. B. Slater, J.S. Rosenblatt, C. T. Snowdon, & M. Milinski (Eds.), Advances in the Study of Behavior (Vol. 24, pp. 47–118). San Diego: A. P. Numan, M., Fleming, A. S., & Levy, F. (2006). Maternal behavior. In J. D. Neill (Ed.), Knobil and Neill's Physiology of Reproduction (pp. 1921–1993). New York: Elsevier. Reisbick, S., Rosenblatt, J. S., & Mayer, A. D. (1975). Decline of maternal behavior in the virgin and lactating rat. Journal of Comparative and Physiological Psychology, 89, 722–732. Rosenblatt, J. S. (1970). Views on the onset and maintenance of maternal behavior in the rat. In L. Aronson, E. Tobach, J. S. Rosenblatt, & D. S. Lehrman (Eds.), Development and Evolution of Behavior: Essays in Memory of T. C. Scheirla (pp. 489–515). San Francisco: Freeman. Rosenblatt, J. S. (1971). Suckling and home orientation in the kitten: A comparative developmental study. In E. Tobach, L. Aronson, & E. Shaw (Eds.), The Biopsychology of Development (pp. 345–410). New York: Academic Press. Rosenblatt, J., S. (1980). Hormonal and nonhormonal regulation of maternal behavior: a theoretical survey. Reprod. Nutr. Dévelop., 20(3B), 791–800. Rosenblatt, J. S. (1983). Olfaction mediated developmental transition in the altricial newborn of selected species of mammals. Developmental Psychobiology, 16, 347–375. Rosenblatt, J. S. (1995). Daniel Sanford Lehrman — June 1, 1919– August 27, 1972. Biographies and Memories of the National Academy of Sciences, 66, 227–245. Rosenblatt, J. S., & Aronson, L. (1958). The influence of experience on the behavioral effects of androgen in prepuberally castrated male cats. Animal Behaviour, 6, 171–182. Rosenblatt, J. S., & Lehrman, D. S. (1963). Maternal behavior in the laboratory rat. In H. L. Rheingold (Ed.), Maternal Behavior in Mammals (pp. 8–57). New York: John Wiley & Sons. Rosenblatt, J. S., Turkewitz, G., & Schneirla, T. C. (1969). Development of home orientation in newly born kittens. Transactions of the New York Academy of Sciences, 31, 23–25. Schneirla, T. C. (1952). A consideration of some conceptual trends in comparative psychology. Psychological Bulletin, 49, 559–597.

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Schneirla, T. C., & Rosenblatt, J. S. (1961). Animal research. I. Behavioral organization and genesis of the social bond in insects and mammals. American Journal of Orthopsychiatry, 31, 223–253. Siegel, H. I., & Rosenblatt, J. S. (1975b). Hormonal basis of hysterectomy-induced maternal behavior during pregnancy in the rat. Hormones and Behavior, 6, 211–222. Siegel, H. I., & Rosenblatt, J. S. (1975c). Progesterone inhibition of estrogen-induced maternal behavior in hysterectomized-ovariectomized virgin rats. Hormones and Behavior, 6, 223–230. Siegel, H. I., & Rosenblatt, J. S. (1980). Hormonal and behavioral aspects of maternal care in the hamster: a review. Neuroscience and Biobehavioral Reviews, 4, 17–26. Silver, R., & Rosenblatt, J. S. (1987). The development of a developmentalist: Daniel S. Lehrman. Developmental Psychobiology, 20, 563–570. Stern, J. M. (1989). Maternal behavior: sensory, hormonal, and neural determination. In F. R. Brush & S. Levine (Eds.), Psychoneuroendocrinology (pp. 105–226). New York: Academic Press. Stern, J. M. (1990). Multisensory regulation of maternal behavior and masculine sexual behavior: a revised view. Neuroscience and Biobehavioral Reviews, 14, 183–200. Stern, J. M., Yu, Y. L., & Crockett, D. P. (2002). Dorsolateral columns of the spinal cord are necessary for both suckling induced neuroendocrine reflexes and the kyphotic nursing posture in lactating rats. Brain Research, 947, 110–121. Terkel, J., & Rosenblatt, J. S. (1968). Maternal behavior induced by maternal blood plasma injected into virgin rats. Journal of Comparative and Physiological Psychology, 65, 479–482. Terkel, J., & Rosenblatt, J. S. (1972). Humoral factors underlying maternal behavior at parturition: Cross transfusion between freely moving rats. Journal of Comparative and Physiological Psychology, 80, 365–371. Williams, C. L., Hall, W. G., & Rosenblatt, J. S. (1980). Changing oral cues in suckling of weaning age rats: possible contributions to weaning. Journal of Comparative Physiology, 94, 472–483. Jay S. Rosenblatt — a close friend Jay was one of my closest friends for nearly 60 years. We first met in the mid-fifties, at a conference in Palo Alto organized by Frank Beach. The members included Jay (with his first wife, Gilda) and Danny Lehrman. Each year one or other of us would cross the Atlantic, and we were in frequent correspondence. The last time I saw him was when he and his second wife, Pat, visited us in Madingley in July 2012. During Jay's last few months we talked by telephone most weekends. Others will be writing about Jay's pioneering work on motheroffspring interaction. There were close conceptual parallels between Danny's work on Ring Doves, Jay's on cats and rats, and my own on canaries, and we helped each other. We viewed development not as an individual matter, but as an ongoing interaction between parent, offspring, and the environment, each changing continuously as a result. But Jay as a scientist had a special quality of persistence in teasing apart the detail of the complex interactions in a developing system. There is always a conflict between seeking for the broad picture and understanding how it all really works, and some became impatient with Jay's careful approach. I recall Frank Beach bursting out in his colorful way, “My J—F—C—Jay, you do not have to tell us that behavior is complicated. That is why we are here!” But it was just that quality that made Jay such a special friend. He would always try to get to the heart of the problem, whether it involved academic or personal matters. He was intensely loyal, and had absolute integrity. I was never sure whether he was happiest painting his sad pictures, working as a Kleinian analyst, or collecting data on maternal behavior.

Once when I asked him, he chose the third, saying he felt more fully himself when doing research. I miss him. Robert Hinde JSR remembered Jay was my official mentor at the IAB, but he was so much more. It all started in 1963 when I began the Master's program in General Experimental Psychology with Jeb Barmack at CCNY. Part way through the first semester, we were discussing my interest in natural behaviors and how well I enjoyed working with the animals in that lab. Jeb knew of the fledgling IAB and recommended that I transfer, which I did in the middle of that first year. At the interview with Jay, I was a nervous wreck and have absolutely no recollections of what we actually discussed. Somehow, Jay saw something and accepted me. This ushered in the very best years of my academic life! Jay was gentle, kind, thorough, patient and most generous. Since I transferred in the middle of the academic year, and the official Rutgers policy was to only accept students in September, Jay arranged for me to begin on his Training Grant as a research assistant to Lorraine Roth. She introduced me to the lab routines and explained her research most willingly, which also added to the positive reputation of Jay as a wonderful mentor. Since I was not officially a student, I could not register for classes, but I did attend the weekly seminars by faculty, students and a very wide range of equally impressive visitors. All of these experiences made indelible impacts and Jay was always there. His insightful questions and comments helped sharpen my thinking about a wide range of subjects. After learning the IAB model of constant age pups to assess changes in parental behavior in adult rats, Jay taught me animal surgery and further observational techniques. As I assisted on a series of investigations of hormonal roles in (what was then called, ‘maternal behavior’) my interest in the role of the newborn in that relationship developed. We discussed this approach and we recognized the problems presented by such a small newborn. Luckily for me, Jay had research experience with felines, and kittens provided me with larger and thus more easily observable newborns. Jay arranged for the creation of a new cat facility in Newark, shortly before the new lab was constructed. Jay worked with me, listened to my proposed needs, added his technical expertise and a new cat facility was in the works. Though initially located in a condemned building, it held a first class colony room (affectionately called, “The Cat House”) and research lab. Eventually we moved to the new lab and we were all under one roof. When Jay published his article on cats in Scientific American he benevolently cited me for my part of one project. The ambiance of the IAB was so positive that faculty, upper level students, new students, undergraduate lab assistants, professional staff and maintenance staff were all treated in a genuinely kindly manner and with full respect. I fondly recall the time Jay drove a few of us somewhere around in Newark in his old DeSoto. While he complained of the flaws in the transmission, he floored the engine in neutral and then dropped it into gear. Here was a living example of the question about proximate and ultimate causation! When I began my permanent teaching career in upstate New York, Jay invited me to visit him at his nearby summer home to discuss final revisions of my thesis. Those were relaxed and productive meetings. Conversations also included sailing his collapsible sailboat, pickled watermelon rinds and the edible flowers that grew by the roadside as well as shopping at the, then new, Sears in Albany. Last year, my wife and I joined Jay and his wife for dinner in New York City. Though we had not met for a number of years, it was just as if we had been in constant contact. His warmth, thoughtfulness, positive recollections of my early career and true personality instantly came through and shall always be a very tender part of my memory.

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In the Jewish tradition, one says about the departed, “May his memory be for a blessing” and here it is certainly true. Thank you Jay. Robert J. Woll Jay Rosenblatt and ethology I once described ethology as “the zoologist's approach to behavior” (Beer, 1963/64). This description would suggest that there are other approaches to behavior besides the zoological. At much the same time, Niko Tinbergen, in a well-known paper in celebration of Konrad Lorenz's 60th birthday, similarly characterized ethology as “the biological study of behavior” (Tinbergen, 1963). He went on to say that the biological approach comprises four different kinds of question: “Huxley likes to speak of “the three major problems of Biology”: that of causation, that of survival value, and that of evolution — to which I should like to add a fourth, that of ontogeny. There is, of course, overlap between the fields covered by these questions, yet I believe with Huxley that it is useful both to distinguish between them and to insist that a comprehensive, coherent science of ethology has to give equal attention to each of them and to their integration” (Tinbergen, 1963). Tinbergen's four questions have since repeatedly structured textbook treatments of the stay of animal behavior. Recently Patrick Bateson and Kevin Laland (2013) reviewed the legacy of these questions in the light of research and theorizing during the intervening fifty years. They argue “that the four-question scheme remains a useful heuristic, but that it requires a more nuanced interpretation than is traditional.” Also they recognize that there are additional questions that are not explicitly comprehended by the original scheme. For instance it is not clear where the questions of behavioral genetics would fit in. With the emergence of behavioral ecology during the 1970s, ethologists became preoccupied with issues of what Bateson and Laland refer to as “current utility” (Tinbergen's “survival value”), to the neglect of questions of mechanism and development. Concern about “inclusive fitness”, “parental investment”, “altruism”, and so on, superseded the wrangles over “energy models of motivation” (Hinde, 1960) and the innate/learned controversy. I recall Marian Dawkins, at the International Ethological Conference in Madison (1986), complaining that people whose business involved dealing with behavioral problems in animals on a daily basis, such as veterinarians and game wardens, were falling back on the discredited motivational theories of “classical” ethology in the absence of help about causal mechanism from current manuals. Bateson and Laland make a plea for a redress of the balance, as Tinbergen would have done. What has this got to do with Jay Rosenblatt? There is no doubt that Jay was a friend of ethology from early on. My first encounter with him may have been at the International Ethological Conference at Cambridge in 1959, when I was in the last stages of my doctoral studies under Tinbergen's direction. He was certainly at the Oxford conference four years previously, for there is photographic proof of it (Fleming, 2006). I was with him at subsequent ethology conferences: Starnberg, Zurich, Stockholm, Rennes, Washington, Edinburgh, Parma, Bielefeld, Madison, Oxford. At the Parma meeting we were both on the Conference Executive Committee. This long engagement with “official” ethology was no doubt favored by his friendship with Robert Hinde, a “second generation” leader among ethologists in Britain. Yet, despite this extended association with the ethological fraternity, it cannot be said that Jay Rosenblatt was a typical ethologist. Unlike Danny Lehrman, his colleague at the Institute of Animal Behavior (IAB) and an avid birdwatcher, Jay showed little if any interest in natural history of the sort that engaged Tinbergen, Baerends, and most of the other European ethologists, when I joined the IAB in 1960 as its first post-doctoral fellow, Jay was only a part-time researcher there. Most days he would leave in the middle of the afternoon to take up his psychoanalytic practice in New York. Prior to qualifying for this occupation

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he had contemplated a career as an artist. Indeed he realized this penchant as an avocation, continuing to paint throughout his life. To the extent to which zoology entered into Jay Rosenblatt's approach to behavior it came from T.C. Schneirla, a comparative psychologist at New York University. Schneirla did remarkable work with army ants, and developed some general ideas about behavioral control and development. His theory of how forced responses of attraction and repulsion in early life become approach and withdrawal under the control of psychological motivation and memory is less well-known than it should be, perhaps in part, because Schneirla's exposition of it is anything but easy reading. However it profoundly influenced a small group associated with the American Museum of Natural History, where Schneirla became Curator in the Department of Animal Behavior in 1943. Jay Rosenblatt belonged to this group. Jay's work at the Museum was on cats, first sexual behavior in males, then the interactions between mothers and their kittens. The sequence of progressive reciprocal influences and transitions in sensory orientation observed in this work set the stage for Jay's subsequent research. On joining Lehrman in his newly founded Institute at Rutgers University in Newark, NJ, in 1958, Jay switched his attention to the maternal behavior of rats. Thus began a collaborative enterprise that went on for 45 years, extending, toward the end, to studies of rabbits and sheep. I leave it to others more conversant with this research than I to spell out the details of the work, remarkable for its originality, ingenuity, meticulousness, and imagination. It impresses me that, at a time when ethology had been largely taken over by concern with issues of “current utility” to the neglect of causal and developmental questions, Jay Rosenblatt was engaged with the latter in a way that shows how they can be integrated in a synthesis that would have met with Tinbergen's requirement. After his retirement Jay embarked on a massive reading survey of the literature on animal parental behavior in search of evidence of evolutionary patterns of descent. The resulting manuscript was unfinished at his death. Nevertheless it helps to make the case that, though starting from a background contrasting with that of traditional ethology, Jay Rosenblatt's approach to behavior included all four of Tinbergen's four questions. Hence it can be argued, as I once did for Danny Lehrman (Beer, 1975), that by a process of convergence, Jay Rosenblatt qualified as an ethologist, by Tinbergen's criteria. A final reflection: Jay Rosenblatt had none of the flamboyance of his once more famous colleague Danny Lehrman. Jay's was a quiet voice, but its echoes may well outlast those of Danny's roar. Jay's students and colleagues not only admired and respected him; they also held him in deep affection (Fleming, Numan, Bridges, 2009). I count myself as one of their number. Colin G. Beer References Bateson, P. and Laland, K.N. 2013. Tinbergen's four questions: an appreciation and an update. Trends in Ecology and Evolution, 28, 712–718. Beer, C.G. 1963/64. Ethology — the zoologist's approach to behaviour. Tuatara 11, 179–187; 12, 16–39. Beer, C.G. 1975. Was Professor Lehrman an ethologist. Anim. Behav, 23, 957–964. Fleming, A. 2006. The three faces of Jay S. Rosenblatt. Developmental Psychobiology, http://dx.doi.org/10.1002/dev.20193. Fleming, A., Numan, M. and Bridges, R.S. 2009. Hormones and Behavior, 55, 484–487. Hinde, R.A. 1060. Energy models of motivation. Sym.Soc.exp.Biol. 14, 199–213. Farewell to Jay Upon completing my undergraduate studies at the Hebrew University, Jerusalem in 1963, following my previous background at an

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agricultural high-school, the study of animal behavior was clearly the direction in which I wanted to continue. However, I was faced with two dilemmas: one, obtaining funding to continue for a PhD; and the other, where could I carry out studies in the field of animal behavior? In Israel my advisors suggested I continue in veterinary studies or animal psychology. But this was not what I was looking for; and I continued reviewing the field of animal behavior. I was seeking a clue that would lead me in the right direction. By chance I finally found exactly what I was looking for in a book, Behaviour of Domestic Animals edited by E.A.E. Hafez in the library of a professor of reproduction at the Weizmann Institute. The book features individual chapters, each dealing with the behavior of a different animal. One of these chapters was written by J. Rosenblatt, and engaged with the behavior of cats. I wrote to Jay and shared with him my ambition to study the subject of animal behavior on the one hand — and my total lack of funding on the other. Jay replied that there was no possibility of a grant for the present academic year, but if I would be prepared to postpone my studies I might be able to apply for a grant for the following year. And so, in 1965, I began my graduate studies at the Institute of Animal Behavior, under Jay's supervision. From the moment that I arrived at the Institute, Jay introduced me to the amazing world of maternal behavior. In my second year there, Jay's work, entitled “Nonhormonal basis of maternal behavior in the rat’, was published in Science, and showed that as a result of long-term cohabitation with pups, virgin rats can display maternal behavior even in the absence of the relevant hormones. Based on the above study and an in-depth analysis of the subject with Jay, it was decided to explore the physiological mechanisms underlying maternal behavior upon parturition by attempting to exchange blood from a rat that had just given birth with that of a naïve (virgin) rat. The aim was to determine whether certain factors in the blood of the mother rat were responsible for the maternal behavior that appears following parturition. The difficulty lay in how to carry out this experiment. How could one exchange blood between two freely-behaving rats in a way that would allow us to observe their behavior? I thus set out to develop a methodology that would enable crosstransfusion of blood between two rats. This took me nearly two years! The technical problems were enormous, from finding suitable cannulae, developing swivels for the cannulae, preventing negative physiological reactions, and cannulae blocked by leucocytes that prevented the free flow of blood, etc. I tried every day, for 18 h daily, to find solutions to the innumerable problems. Throughout all this frustrating period Jay's encouragement and support were unending, both financially in regard to the costs of time and equipment for experimentation, and emotionally. Without doubt, had it not been for Jay's untiring patience and reassuring presence during this very long period, I would have given up. In addition to his being an external partner to the research through funding and moral support, Jay was a highly practical work partner too, and after we had worked out the surgical technique of implanting the cannulas and exchanging blood, Jay took an active part in the experiment itself. Despite Jay's lesser experience in micro-surgery, his motivation to perform every stage of the procedure was astounding. Undoubtedly, the practical experience contributed to him when he wrote up the article for me that described the technique (Terkel, 1972): “A chronic cross-transfusion technique in freely-behaving rats using a single heart catheter. “Even today I still appreciate this gesture on his part.” The results of the study (Terkel & Rosenblatt, 1972) showed that naïve female rats receiving blood from parturient females became maternal, and this opened up a fertile field of investigation of the hormonal control of maternal behavior in the rat. Aside from Jay being a scientist with broad horizons, able to focus on the most important and relevant questions in the field of hormones and behavior and specifically issues of maternal behavior, he was also a warm human being, sensitive and concerned. Upon my arrival in the

U.S. in 1965, I could not have wished for a more wonderful person, as both mentor and friend. One of the moments that I will always remember took place in June 1967, with the outbreak of the Six Day War, when I decided to return to Israel. Danny Lehrman called me into his room, where Jay was waiting. With hugs and encouragement, they promised that whenever I was ready to return everything would be waiting for me as if I had never left; and they wished me a safe journey. The immense feeling of warmth and friendship in that room cannot be described… In 1971 I was accepted as a lecturer in the Department of Zoology at Tel Aviv University. Since then, I have had the good fortune to mentor and supervise a great number of MSc. and PhD students. To each and every one of them I have sought to provide a scientific and welcoming atmosphere, similar to that which I received from Jay. At times I have felt that I was continuing the positive tradition with which I had been ‘imprinted’ and inspired by Jay during the five years that I spent at the Institute of Animal Behavior during my doctoral studies. And Jay — I shall forever be grateful! Joseph Terkel References Rosenblatt J S and Schneirla T C 1962 The behaviour of the domestic cat pp 388–453, in Hafez ESE editors. The Behaviour of Domestic Animals. Cambridge: Baillière, Tindall & Cox, Ltd.; 1962; Rosenblatt, Jay S. (1967). “Nonhormonal basis of maternal behavior in the rat”. Science 156 (no. 2781): 1512–1513. http://dx.doi.org/10. 1126/science.156.3781.1512 Terkel J 1972. A chronic cross-transfusion technique in freely behaving rats by use of a single heart catheter jap.physiology.org/content/jap/33/4/ 519.full.pdf Terkel, Joseph; Rosenblatt, Jay S. Humoral factors underlying maternal behavior at parturition: Cross transfusion between freely moving rats. Journal of Comparative and Physiological Psychology, Vol 80(3), Sep 1972, 365–371. http://dx.doi.org/10.1037/h00329651 Jay S. Rosenblatt — pioneer in parental behavior research, mentor, friend Jay, our mentor, had a rare quality — he was nurturant to his core. His research on parental behavior and development was a coherent extension of his humanistic quality as a painter and psychotherapist. Jay's combination of being creative, respectful, non-judgmental, generous, and pragmatic gave him the uncommon virtue of making those with whom he interacted feel that they were part of the solution, rather than of the problem. My connection with Jay: Barry Jay was the person who not only opened the door for me to enter the field of animal behavior research, but he also made it possible for me to continue in my career at crucial turning points. My first contact with Jay was when I was just a seven years old kid in the summer colony in Goldensbridge, in upstate New York, where Jay and his lovely wife Gilda stayed with her sister in her home and my parents also had a summer home. I got to know Jay every summer leading to my teenage years. Jay recruited my best friend, Peter Gold, to be a research assistant in the Department of Animal Behavior, which is on the 5th floor of the American Museum of Natural History in Manhattan, where Jay was performing his doctoral dissertation research on cat sexual and maternal behavior. When Pete got a well-paying job, in the summer that I was 16, he asked me to replace him at the Museum. I took the opportunity enthusiastically. Through that connection, in my senior year at City College, I took a course with Danny Lehrman, who was a visiting professor there from Rutgers–Newark. Danny invited me to be his doctoral student, so I applied and got accepted to Rutgers University with Danny as my mentor 1

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at his newly-founded (in 1958) Institute of Animal Behavior (IAB). Jay had joined Danny at the Institute by the time I arrived and he became my “second mentor”. After I defended my dissertation, I was accepted for a postdoc by Charles H. Sawyer at the Brain Research Institute at UCLA. When I joined his laboratory, he was recording neuronal responses correlated with the pseudopregnancy reflex (which is elicited by vaginocervical stimulation) in rats. Danny, with Jay's approval, offered me a position on the IAB faculty, and I started in 1966. I was fortunate in being permitted full freedom to develop my own line of research. Through a fortunate invitation to spend a summer in James Olds' laboratory at the University of Michigan, I first observed the lordosisinducing/immobilization-producing effect of vaginocervical stimulation in awake, freely-moving rats and subsequently, with Knut Larsson and Josh Wallman, we found that vaginocervical stimulation blocked behavioral and single-neuronal responses to painful stimulation. All of us in the IAB were devastated when Danny died suddenly of congestive heart failure in the summer of 1972. As soon as we faculty started to recover from the shock of losing Danny, we met as a group and, needing virtually no discussion, by unanimous acclamation, elected Jay to take over as Director of the Institute. Jay was crucially supportive to us faculty and the students of the Institute. He applied for, and was awarded, an NIH Training Grant, which continued for many years, providing support for doctoral and postdoctoral students, the lifeblood of the Institute. When I was having trouble getting grant support for my reproductive behavior research, Jay bailed me out financially for several years, providing the funding for my technician, Cindy Banas. When I endeavored to identify the neurotransmitter(s) released into the spinal cord by vaginocervical stimulation that trigger the analgesia, Jay negotiated a special, remarkably generous, funding allocation via the Rutgers University Vice President, which enabled me to collaborate with Frank Jordan of our Chemistry Department, which supported our further research and led to our patent for an analgesia-producing peptide. Having found suggestive evidence in rats that vaginocervical stimulation blocks pain, I got to a point in my research when I decided that the only way of knowing for sure whether vaginocervical stimulation blocks pain is to ask women. This led to my first human research study. I obtained IRB approval to study the effect of vaginal or cervical selfstimulation in women. That was the precise point at which Jay played another crucial role in my career development. Some IAB faculty members made a representation to Jay, in his position as IAB Director, to prevent me from performing research on human sexual behavior in the IAB, because “it would give the Institute a bad name”. He flatly rejected their request. Jay staunchly defended my academic freedom to perform human research in the IAB. My connection with Jay: Cruz From 1978 to 1980 I was an undergraduate student in the Autonomous University of Madrid (UAM), assisting J.A.I. Carrobles in his research on learning in rats, which he was performing in Prof. Antonio Guillamon's laboratory at the School of Medicine. Guillamon and his student, Santiago Segovia, were studying the morphology of the accessory olfactory and vomeronasal system structures (AOS/VNS) in relation to the effect of perinatal gonadal hormones on sexual differentiation of the brain. Guillamon suggested that for my senior research thesis I study sex differences in learning, and I decided to focus on the partial reinforcement effect. On the basis of this research and my grades, I was contracted by UNED (Universidad Nacional de Educacion a Distancia) as “Profesora Colaboradora”, in Psychobiology, which was an entry-level faculty position. This involved teaching, my own doctoral research, and assisting Guillamon in his research. After several collaborative publications that demonstrated sexual dimorphism in the VNS, which was based solely on neuromorphological and endocrinological evidence, I raised the question of whether there is a specific behavioral role of the sex differences in the VNS in the known dimorphism of reproductive behavior.

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Because I was already familiar with Jay Rosenblatt's research, I chose to study maternal behavior. Over the next five years, I developed my own line of research and a research team on maternal behavior, which enabled me to gain a tenured position in the Psychobiology Department of UNED. Our studies showed that lesioning of VNS structures via different procedures (electrolytic, chemical, and excitotoxic [of GABAA receptors]) induced maternal behavior in male rats, suggesting that the male VNS normally inhibits maternal behavior. During this time, I developed a computer program to record maternal behavior sequences, which was a significant advance in the recording of maternal behavior. I wanted to improve my methodological skills in the field, and as Jay was the world's foremost investigator of maternal behavior, I applied for, and was awarded an international visiting professorship to work with Jay. I arrived at the IAB as a visiting professor in May, 1990. Some months before starting at Rutgers, Jay visited me in Madrid on his trip to Goteborg, Sweden, in order to receive the Honoris Causa Doctorate at the Swedish University, a major scientific academic acknowledgment. Upon getting to know Jay better over the years, I believe that Jay's first visit to Spain was based on his prudence, preferring to see, first hand, my laboratory and my way of dealing with research before welcoming me to Rutgers. Jay arrived with Carlos Beyer to our psychobiology laboratory in the National University of Distance Education (UNED) in Madrid. They each presented several interesting seminars and interacted with our research group. This was the beginning of a long close friendship and fruitful scientific collaboration with both Jay and Carlos. This unforgettable personal and professional interaction resulted in multiple publications and in an UNED Honoris Causa Doctorate, which made Jay part of our academic community in 1997. Just a few months before he passed away, as a further acknowledgment of his international inspirational influence in behavioral neuroscience, Jay received the Gold Medal of the Faculty of Psychology, UNED, in November, 2013, which I had the honor of presenting it to him in person at his home in New Jersey (Fig. 1). It is painful for me to write these memories as both Jay and Carlos are no longer with us. But it is an honor and I feel grateful to have this opportunity to express how important in my professional activity and life was Jay as a scientific and personal role model. Our collaboration with Jay In discussion with Jay we decided to map the brain regions involved in the expression of maternal behavior. One of our doctoral students, Byron Johnson, had recently employed 2-deoxy-glucose autoradiography in his dissertation, so we applied this methodology to the maternal behavior study. Jay guided a complex and insightful experimental design to compare maternal behavior induced by a variety of hormonal and environmental conditions, i.e., natural pregnancy-induced, hysterectomyinduced, and pup-induced in virgin females. Designing the study was an exciting, sometimes stressful, process of negotiation among five researchers of different scientific traditions of three different countries ranging from unmanageable complexity to naïve simplicity that Jay managed to resolve in his characteristic calm, respectful, unflappable, psychotherapeutic, scientifically rigorous, style. Thanks to Jay, and his quietly efficient, decades long-term loyal research associate, Anne Mayer, everybody ended up happy with the final experimental design, and the end result was that the findings of this study have been cited in psychobiology textbooks. In the results of this research, Jay recognized an interesting interaction between hormonal condition and behavioral expression such as activation of several vomeronasal regions in virgin versus intact mothers. The question arose as to whether regions whose activity was lower than baseline were simply deactivated or whether they were actively inhibited, and if the latter, which regions were generating the active inhibition. To answer this question, in parturient, sensitized, and hysterectomized rats, we combined the 2-DG and c-fos methods on the basis that the 2-DG method shows activity in the presynaptic terminals, while the

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c-fos method shows activity in the postsynaptic cell bodies. We found that in some regions (e.g., medial preoptic area) there was both 2-DG and c-fos activation, but in others, (e.g., accessory olfactory bulb), 2-DG activity increased whereas c-fos activity decreased. The former is evidence of active excitation, whereas the latter is evidence of active inhibition. These findings were gratifying to Jay because they enhanced his previous studies, based on which he developed his concept of the importance of olfactory input to the onset of maternal behavior under various hormonal and stimulus conditions. They also supported his non-hormonal, sensory-dependent concept of the control of postpartum maternal behavior. We admire Jay's remarkable mind and indefatigable intellectual vigor and rigor. This made Jay a powerful role model for all of us to follow in his style of being. For years, every year, he threatened to retire “next year” and relax with his delightful loving wife, Pat; fortunately for his students and colleagues, he didn't. But even after he finally did retire, he continued to come into his office several days a week every week to work on his review of the evolution of parental behavior. We had many conversations about this topic. Moreover, Jay made perceptive recommendations about our own research, which contributed significantly to improve our experimental designs and focus our research questions, which we are following up on even today. Remembrance We remember as one of the most characteristic features of Jay's wellrounded personality was that he was able to move comfortably and spontaneously from his fond memories of his deceased first wife Gilda, his loving and proud feelings about their children, Nina and Danny and granddaughters, to relevant and salient issues on our field and our research on which he was always current on the literature. We are not exaggerating when we conclude that one of Jay's most important lasting contributions to our field is his mentorship, encouragement and supportiveness, which played a major role in launching the scientific careers of young investigators not only throughout the USA but also in Mexico, Canada, Spain, France, Sweden, Germany, Italy, England, Russia, and Brazil. By imprinting the importance of parental behavior on so many young investigators who came in contact with him, Jay is truly the father of the School of Parental Behavior Research worldwide. Maria Cruz Rodriguez del Cerro Barry R. Komisaruk

1984: Jay and Barry sharing a light moment at Jay's 60th birthday reunion conference at the IAB.

1984: Jay's research group at his 60th birthday reunion conference.

Rosenblatt memorial

Fig. 1. November, 2013: Cruz has just brought to Jay his award of the gold medal of Honoris Causa Doctorate from UNED.

I had three thesis advisors: Daniel Lehrman, Jay Rosenblatt, and Colin Beer. Danny was inspiring, Colin fed my appetite for philosophy and the history of science. But Jay was my teacher and mentor. Jay taught me how to write. When Danny died in late August, 1972, I had finished my thesis work and was beginning to write up the thesis. Jay stepped in at that point and, in addition to holding the Institute together, he taught me the nuts and bolts of how to write science. I would write a section, give it to Jay and, within a few days, he would give it back absolutely covered with red ink. Each was a constructive comment showing what, where or why my writing was confusing. In our discussions he impressed upon me that he was not suggesting that I use his wording or structure. Rather, his comments were designed to show me that what I had written was confusing or downright sucked and I had to change it. So while Danny's writing was elegant, there was simply no way (in my opinion) that someone like me could get from where I was to that level. Jay taught me the elements and the way in his own patient way. I took this lesson to heart and I believe anyone who has written papers with me knows that I have rapid turn-around, that my comments are more

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a flag for places for improvement, and that my revisions are suggestions, not dictates. Many have also seen me haul out my thesis to illustrate Jay's comments. My two favorite courses at the Institute were Jay Rosenblatt's class in social development and Colin Beer's ethology class. This was because each taught from an historical perspective. I have always loved the history of ideas, and both Jay and Colin spent quite a bit of time showing how the foundation of the questions being posed at that time was rooted in philosophy and biology. I have kept this interest and teach an occasional course entitled “Before there was PubMed”. The motivation is the common complaint of ‘old farts’ (now us) that the ‘new turks’ (then us) do not read the original literature (or even the literature that was current when we were graduate students). Sometimes this comes from an unwillingness to leave the lab to go to the library and reading papers that might have been published in the 1980s; certainly never papers from the late 1800s to early 1900s. I contend that most, if not all, questions in biology were considered in depth back then but, because of politics or fashion, fell out of favor and were forgotten. Thus, much of what is the state-of-the-art is actually technique driven, often meaning that the question has had to be rediscovered. It is useful to recall that ‘ancient’ scientists (however one wishes to define that) thought deeply about the essential problems and were not distracted like present scientists tend to be. Jay launched the research academic careers of many graduate students, and fostered the development of postdoctoral fellows, who now have attained prestigious academic positions. Jay instilled a passion and excitement for science into those fortunate enough to work with him because he was passionate and excited about his own science. David Crews Memories and appreciation As I look back on my friendship with Jay, I realize that it didn't arise in the usual way among academics: for example, during membership on the faculty or the student body of the same institution, or from being a student or (far less likely) a teacher of his. Nor did my friendship arise from growing up with him or in some shared activity outside our professions. Jay and I came together over the years through a common interest — perhaps even a passion — for studying the early development of infants within the mother–infant relationship. More concretely, I met Jay in 1967, through the Department of Animal Behavior that existed then at the American Museum of Natural History, and through Dr. Ethel Tobach in particular. I was doing a late postdoctoral fellowship at the Museum, with Ethel, as part of an NIMH program to bring young psychiatrists into biological research. Jay and Daniel Lehrman had left the Museum a few years previously to start their new department at Rutgers, but Ethel directed me toward them and toward Jay's research on kittens and their mothers in particular. I think it was Jay's approach to studying how kittens develop the ability to find their mother's nipples and attach to them, that inspired and taught me how to design a series of experiments that analyze, in steps, the roles of different sensory systems and simple behaviors in the sequences of learning and adaptation that develop within the apparently simple interactions that we call a mother–infant ‘relationship’. I remember visiting Jay's lab at Rutgers from my lab in the department of psychiatry at Albert Einstein School of Medicine, many times over the ensuing years, talking to Jay about his work, occasionally presenting my work at his lab meetings and importantly, meeting a number of his postdoctoral students including Christina Williams, Ted Hall and a number of others. But it is not my visits to Rutgers in those years that I remember most distinctly, but rather my talks with Jay over many years at the annual meetings of the Society for Developmental Psychobiology, meetings that Jay and I rarely missed. There were years that Jay hardly missed a

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presentation at those meetings, and in my memory at least, he always wore his plain red suspenders. There were some years when he seemed to raise a point for discussion after every single presentation; and Jay's points were never trivial. In fact they often lead to fields seemingly distant from the subject of the talk, to which he almost always made an interesting connection. At these meetings we often went out to supper together with other members of the society and it was here that I learned about Jay's watercolors with which he seemed as involved as his research. I also found that Jay and I shared another occupation: as psychotherapists. I always imagined Jay as a particularly good psychotherapist because of his patience and his intuition — both of which are handy as well for doing research. In later years, I found myself on an NIMH research committee site visit on Jay's program at Rutgers. This experience made me realize how much I still thought of myself as Jay's student, and particularly when I made a somewhat awkward defense of Jay's approach in an argument I had with another committee member, Frank Beach, an even more August figure than Jay himself at that time. Over the more than 45 years that I knew Jay, I came to realize and appreciate the affection I developed for him, even though we got together rarely and at considerable intervals. Now that he is gone, I realize even more clearly what a very good friend he has been in my life. Myron Hofer Remembering Jay Rosenblatt It was early spring, 1973. I just finished by doctoral research at the University of Chicago and was writing up my dissertation. At that time, my wife, Marilyn, and I had three children. Therefore, I was not initially considering a postdoctoral research position, but instead was sending out my rather skimpy curriculum vitae to all universities and colleges with tenure track job openings in behavioral neuroscience (physiological psychology). Fast forward to the summer of 1973: although I had some nibbles, a job did not materialize. I was getting very desperate. Fortunately, at the end of June, the Eastern Conference on Reproductive Behavior (a predecessor of the Society for Behavioral Neuroendocrinology) was held at the University of Chicago, hosted by my thesis advisor, Howard Moltz. It was at this conference that I presented my doctoral dissertation research, and Jay Rosenblatt was in the audience. Subsequently during the conference, Howard and I approached Jay and asked whether a postdoctoral position would be available for me at the famous Institute of Animal Behavior (IAB) at Rutgers University. Jay indicated that he would get back to us as soon as possible. As it turned out, Jay and Barry Komisaruk each came up with $4000 from their grant sources, which allowed me one year as a postdoctoral researcher at IAB with a stipend of $8000. During this first year, I applied for an NIH Postdoctoral Fellowship, which funded an additional year, and I spent two wonderful years at IAB, where Jay was the Director. Other postdoctoral opportunities eventually became available to me during the summer of 1973, but I accepted the offer of Jay and Barry, which began my 41 years of friendship with Jay. One of the important reasons for working with Jay, in comparison to the other opportunities that I had, was that postdoctoral research under Jay's mentorship allowed me to continue my research on the brain mechanisms regulating maternal behavior in rats. I wanted to remain in the maternal behavior field, and Jay was a leader in the field. At Chicago, I found that the medial preoptic area (MPOA) was essential for maternal behavior in postpartum rats. At IAB, based on the research of Siegel and Rosenblatt (1975) that estradiol was critical for the onset of maternal behavior at parturition, Numan, Rosenblatt, and Komisaruk (1977) showed that estradiol acts on MPOA to stimulate maternal behavior. In doing this research at IAB, Jay allowed me a very high level of independence, and that prepared me well for my subsequent academic career. Life at IAB under Jay's directorship was an absolutely incredible scholarly experience. The Institute of Animal Behavior was a research

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institute consisting of a group of faculty, graduate students, and postdoctoral fellows who were studying different aspects of animal behavior: (1) ethology and evolution of behavior; (2) developmental processes; (3) neurobiology of behavior. The behaviors that were studied were natural behaviors, and the species examined included rhesus monkeys, rats, ring doves, sea gulls, and lizards. As a demonstration of the tremendous scope of the research that was conducted at, or influenced by, IAB during Jay's directorship, a volume of the Annals of the New York Academy of Sciences documented many aspects of IAB's broad research program (Komisaruk, Siegel, Cheng, & Feder, 1986). Although I am primarily a neurobiologist, being immersed in this environment had a tremendous influence on promoting my interests in understanding how evolutionary forces, genetics, and developmental processes affect the neural circuits that control social behavior. Although most researchers at IAB conducted their work within the boundaries of their primary focus (ethology, development, or neurobiology and neuroendocrinology), because of the diversity of ideas at IAB, at times a wonderful amalgam was created, and the prime example that comes to my mind is the research of David Crews. Jay Rosenblatt and his students created our modern understanding of the biology of maternal behavior, and an overview of Jay's important framework and formulations has been written (Fleming, Numan, & Bridges, 2009). I would like to focus on two of Jay's early works that set the stage for much of the subsequent research that has been conducted on the role of hormones and brain mechanisms in the control of maternal behavior. In one of the best chapters on maternal behavior in rats (Rosenblatt & Lehrman, 1963), Jay reported (1) that the onset of immediate maternal behavior is triggered by the physiological events associated with parturition and (2) that if a mother rat is not allowed to interact with her young near the time of parturition, maternal motivation and interest in young declines. In a subsequent study, Jay defined the characteristics of pup-stimulated maternal behavior, or the sensitization of maternal behavior in virgin (nulliparous) rats (Rosenblatt, 1967). Although virgin rats that are not exposed to the physiological events of parturition do not show immediate maternal behavior, if such rats are continuously exposed to freshly nourished pups over several days, then after about seven days of such exposure, the virgins begin to act maternally toward the young. Additionally, in this study, Jay showed that such pup-stimulated maternal behavior has a nonhormonal basis, as it occurred in hypophysectomized and ovariectomized virgin rats. All of these findings indicated that there is a basic neural circuitry that is operative to promote maternal behavior in nulliparous rats, that physiological factors associated with parturition act on this circuitry to stimulate immediate maternal behavior in primiparous rats, and that in order for maternal behavior to be maintained during the postpartum period, the mother must interact with her young near the time of parturition. These significant early observations led to important questions that provided direction for much of the subsequent research on maternal behavior. (1) What are the specific physiological and hormonal events that trigger immediate maternal behavior at parturition? (2) How do hormones act on brain mechanisms to shift the longlatency maternal behavior shown by virgins after pup exposure to the immediate maternal behavior shown by primiparous females? (3) What are the mechanisms that allow maternal experience with pups at parturition to cement in place the mother–infant bond so that maternal behavior can continue during the remainder of the postpartum period? Answers to aspects of these questions, which affirm the significant impact of Jay's initial research, can be found in several reviews (Bridges, 2008; Numan, Fleming, & Levy, 2006). Importantly, many of the answers to these questions resulted from the research of Jay and his students. Jay was not only interested in the biological mechanisms of maternal behavior, but he also did research on the development of behavior (see Fleming, 2007). Further, Jay was interested in the evolution of behavior, and in the last years of his life he was writing a manuscript on the ‘Evolution of Parental Behavior among the Vertebrates: A Review’. Just before he died, he sent me a copy of this unfinished manuscript, which I

will always cherish. In addition to being an outstanding scientist, Jay was a kind, gentle, and considerate person. Marilyn and I will miss him very much. Michael Numan References Bridges RS (ed.). (2008). The Neurobiology of the Parental Brain. San Diego, CA: Elsevier. Fleming AS. (2007). The three faces of Jay S. Rosenblatt. Devel Psychobiol, 49, 2–11. Fleming AS, Numan, M, Bridges, RS (2009). Father of mothering: Jay S. Rosenblatt. Horm Behav, 55, 484–487. Komisaruk BR, Siegel HI, Cheng MF, Feder HH (eds.). (1986). Reproduction: a behavioral and neuroendocrine perspective. Ann NY Acad Sci, Volume 474. Numan M, Fleming AS, Levy F. (2006) Maternal behavior. In: Neill JD, ed. Physiology of reproduction, volume 2. San Diego: Academic Press, pp. 1921–1993 Numan M, Rosenblatt JS, Komisaruk BR (1977). Medial preoptic area and onset of maternal behavior in the rat. J Comp Physiol Psychol, 91,146–164 Rosenblatt JS (1967). Nonhormonal basis of maternal behavior in the rat. Science, 156, 1512–1514. Rosenblatt JS, Lehrman DS (1963). Maternal behavior of the laboratory rat. In: Rheingold HL, ed. Maternal behavior in mammals. New York: Wiley, pp. 8–57. Siegel HI, Rosenblatt JS (1975). Hormonal basis of hysterectomyinduced maternal behavior during pregnancy in the rat. Horm Behav, 6, 211–222. My reflection on Jay S. Rosenblatt I first met Jay in the summer of 1975 when I joined the Institute of Animal Behavior as a post-doctoral trainee with Harvey Feder. I had received my doctoral degree at the University of Connecticut from Drs. Mike Zarrow, Vic Denenberg, and Bruce Goldman where my thesis focused on the endocrine regulation of maternal behavior in the rat. In hindsight Jay should have been my postdoctoral mentor given our mutual interests in the regulation of maternal care. However, in fact, he became my intellectual mentor as I pursued my research training at the IAB. During that period, we had an active difference in opinion regarding the relative role of estrogens in the onset of maternal behavior. As a skeptical postdoctoral fellow, I questioned the existing emphasis on the role of estradiol in this process. I recall that after completion of one study that examined the role pregnancy termination on the onset of maternal care, we ended up interpreting the findings somewhat differently. The temporal necessity for estradiol after surgery in the pregnant rat for the rapid onset of maternal behavior came into question. This made for a very interesting back and forth for the manuscript discussion. Not one to easily give in, I was a bit headstrong and became frustrated to the point that I decided to submit the paper emphasizing my perspective. Of course, this came to light, and I'm sure frustrated Jay. However, given his openness and temperament, he never chastised me, but rather we sat down and came to an acceptable agreement on what was to be our publishable version. Over the years, I came to really appreciate his mentorship, although this never was talked about much in our interactions. Jay had a wonderful intellectual and humane persona. I have many fond memories of Jay, one notably one being his joining students, postdocs and staff in the IAB conference room for lunch. Wearing suspenders and somewhat baggy pants, Jay would pull out his pocket knife and proceed to peel an apple and cut some cheese in what would become an intriguing discussion about science, politics, art or music. Jay's interests and accomplishments extended beyond the field

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of science into the arts where we were an accomplished painter and into the humanities where he was a counselor. Jay's trainees in the area of maternal behavior included an impressive set of students and postdoctoral fellows. These included Joe Terkel whose studies using parabiotic blood transfusions provided evidence for the humoral mediation of maternal behavior in prepartum rats. Likewise, Alison Fleming shortly thereafter generated seminal research on the role of the olfactory system in maternal behavior, Harold Siegel demonstrated a role for estrogen in pregnancy-terminated rats in maternal care, and Michael Numan reported that the medial preoptic area was a site of estrogenic stimulation of maternal behavior. It was an honor to continue this line of behavioral research as a postdoctoral fellow with Jay and Harvey in the late 1970s. As a postdoctoral fellow, I gained an awareness and appreciation of viewing behavior from an ethological perspective, a perspective that Jay developed in association with Schneirla and Lehrman. For me this perspective has been a major influence in my career in my efforts to understand normative physiological and neural processes that influence the expression of maternal care over the course of development and the animal's lifespan. Perhaps Jay's most significant research contribution to the field of maternal behavior was his 1967 Science paper on the “Nonhormonal Basis of Maternal Behavior in the Rat.” This paper established that the neural capacity of the female to respond maternally was present in all females. This was to serve as the foundation for subsequent research that examined the stimulatory role of hormones and neuropeptides in the biochemical regulation of maternal care. Although published more than 50 years ago, it still is cited as a seminal publication. Finally, personally I was happy in recent years to have developed a closer personal friendship with Jay. We corresponded the past few years, sharing both professional and personal experiences. I certainly will miss him as both a friend and colleague. His positive impact on science and our acquaintance is lasting. Bob Bridges Thinking of Jay When I think of Jay the first picture that enters my mind is a calm face with a faint smile and two blue eyes looking deeply at me (into me). He inspired trust and awe at the same time: a deep thinker and a good listener, too. I was neither his student nor his post-doc; rather, I think of myself as Jay's scientific grandchild. My relationship with him stems from his endless scientific curiosity: Jay had been wanting to test his idea that, after parturition, the maintenance of maternal behavior depended not so much on hormones but on the contact between mother and young. He had already obtained evidence in rats to support this hypothesis but, as these mammals interact with the litter for many hours a day, he wanted to find out if shorter contact with the newborn would suffice to maintain maternal responsiveness. Rabbits were the ideal species to test this because they nurse the kits only once a day for 3 min. The problem was Jay had no access to rabbits! So he contacted my mentor (Carlos Beyer), who had done much work on lactation in rabbits and whom he had known for many years. He had established a new research center in Tlaxcala (central Mexico), supported by the Center for Research and Advanced Studies of the National Polytechnic Institute (CINVESTAV) and the Autonomous University of Tlaxcala, and the facilities included a large rabbit vivarium. Jay and Carlos agreed that he would come to Tlaxcala to discuss the possibility of starting a line of research on rabbit maternal behavior. I had never worked on maternal behavior nor was I particularly interested in the topic: I was happily investigating sex behavior in rats for my PhD dissertation. But Jay came to our lab and we started talking and he presented to me a whole new area of Behavioral Neuroendocrinology that I hardly knew. He emphasized that the investigation of rabbit maternal behavior had been stopped after the untimely death of Mike Zarrow, a pioneer of such work. He highlighted that, by having a good rabbit vivarium, we were in a unique position to re-start the investigation of the neuroendocrine

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regulation of maternal behavior in such a mammal. Carlos Beyer was enthusiastic at this proposal as his original work in rabbits (with Flavio Mena) had focused on the neuroendocrinology of lactation, not on behavior. Jay's visit occurred in 1989, when we started planning the first experiment to describe and quantify nest-building and to measure the associated variations in specific hormones in blood across pregnancy. We presented the first results in 1991 at the meeting of the Mexican Society of Physiological Sciences. In 1994, Jay invited me (and Fréderic Levy and Benoist Schaal) to give a talk in the symposium on parental behavior he was organizing for the Winter Conference on Current Issues in Developmental Psychobiology (in the island of St. Lucia). This was my first invitation to lecture at an international meeting and it exemplifies one of Jay's many qualities: his support of young investigators and his trust that they will develop along new, original paths. This indeed occurred as the two afore-mentioned French researchers are now well recognized figures in the areas of sheep maternal behavior (Levy) and mammalian pheromones (Schaal). My own work on rabbits started with our first joint publication in Physiology and Behavior in 1994, largely supported by a grant from NIH that we renewed three times. Our last joint publication, in 2005, was not the end of our relationship, neither in scientific nor in personal terms. Jay and I shared many extra-scientific interests and we enjoyed each other's company. Across more than 12 years we would see each other once or twice a year at a scientific meeting, in the US, Europe or Mexico. On many of these occasions he would be joined by his wife Gilda and then, they spoiled me. I remember him driving through back roads (looking for the garage sales he loved) or on the suburban highways (looking for the outlet malls I loved). His appreciation of art of all kinds allowed Carlos Beyer and me to take him to unusual places in Mexico (like Oaxaca) to look at the baroque churches, buy the handmade rugs, or eat grasshoppers! Yes, Jay showed his “joie de vivre” in many forms like when, (in his seventies) shortly after his marriage to his second wife Pat, he told me very excitedly that they'd just bought a twenty-year share-time condo in Puerto Vallarta! I was able to say good-bye to him on the phone, a few days before he died. I told him I loved him very much and that I would not be the scientist I am without the support, affection, and trust he had given me across many years. Jay replied: “That's nice to hear.” Gabriela González-Mariscal Remembrances of Jay S. Rosenblatt by a reluctant daughter It is incomprehensible to me that although I have known Jay Rosenblatt for well over three decades, my mind conjures up our discussions, and the sight of him, during the last days of his life. My husband and I had traveled out to Hackensack to see Jay during one of the impressive snowstorms of winter 2014. Jay was lying in bed, attached to an oxygen tank, having trouble talking. He looked at me and said: “You look so tired, Rae — you should rest”. That tells the tale of my relationship with Jay. He was not my supervisor, nor my mentor, nor my teacher. The famous father of mothering was a “mother” to me. He offered advice on all matters of life and addressed his clinical interviewing expertise to investigating me and my relationships. Like a good child, I fought most of his efforts. Jay was at peace with himself on those last few days. He had organized his worldly affairs, arranged things for his family, had considered all his complex emotions about living and dying, and was ready. He said that he did everything he was told to do, and everything he thought of himself. Jay's unique aura came from his quiet, calm bearing and manner. One always had the feeling of being in the presence of a psychoanalyst, and so it was toward the end. For Jay it seemed that all questions had complex answers, and all problems rested on a long history that started before birth, but could be unraveled or deconstructed, with endless patience and analysis. In Fleming's 2007 review of Jay's impact on the study of maternal behavior, she lists as his major contribution to his field, two ideas that reflect his perspective. The first idea is that learning can occur at very early ages,

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and can be demonstrated by using appropriate ecological and sensory conditions. The second is the concept of behavioral transitions in maternal behavior and behavioral synchrony between mother and young. Each of these perspectives emerges from Jay's mix of creativity, psychoanalytic perspective and experimental research. Like all superb ideas, these have become so integrated into thinking that they now appear self-evident. Upon reflection, Jay was happy with this, and he was thrilled with Fleming's analysis. My own research has focused on the circadian timing system (reviewed in Butler et al., 2009). This work deals with the observation that most physiology and behavior of mammalian organisms have daily oscillations. These rhythmic processes are orchestrated by an internal circadian timing system whose phase is set by signals from the external environment. In mammals, the circadian timekeeping system is composed of a central pacemaker in the brain's suprachiasmatic nuclei (SCN) and subsidiary clocks in virtually every cell of the body. The SCN is synchronized to local time mainly via photic cues detected by the retina. The activity of the SCN results in circadian fluctuations in physiology and behavior. SCN output is communicated via neuronal efferents and diffusible humoral cues that reach local brain regions and are transmitted thence to the rest of the body. As someone more removed from the experimental work on maternal behavior, I see Jay's trajectory thoughtfully delineated in the reviews that he wrote. These reviews point to principles of organization that apply not only in the study of maternal behavior but also in other realms of research inquiry. Each reflects the ideas and tools available for analysis over the decades that he worked. As each new complex of ideas and tools emerged over the decades, Jay incorporated these into his thinking about maternal care and parent–young interactions. The following paragraphs point to his trajectory as I tracked it and the parallels in my own field of study. One of Jay's first reviews involved a focus on hormonal and nonhormonal regulation of behavior (Rosenblatt 1980). The integration of hormonal and nonhormonal factors in guiding behavior and physiology is a continuing theme in all manner of research. A small representative example from work in my laboratory is our demonstration that the subset of cells in the suprachiasmatic nucleus that express androgen receptors also receive direct photic input from the retina via the retinohypothalamic tract (Karatsoreos et al., 2011). These cells integrate endogenous and exogenous information to set the phase of the brain clock. From integration of hormonal and non-hormonal factors, Jay turned to an examination of what Fleming termed “transitions” and I will generalize to “anticipations”. How does a previous endocrine or stimulus situation prepare the animal or human for what is about to occur next. This is exemplified in Jay's review of how the hormones of pregnancy support the onset of maternal behavior in the rat (Rosenblatt et al., 1988). The transitions from one state to another are key to understanding the organization of behavior. The principle of preparing for change in hormonal and behavioral transitions is not only at the heart of hormones and maternal behavior but also understanding the circadian timing system. The principle of transitions/anticipations is really the essence of the study of circadian rhythms. The brain clock sets the phase of oscillators throughout the body, thereby preparing the organism for events that are about to occur. One key example is seen in the anticipation of seasonal cycles wherein the reproductive system anticipates the spring — optimizing the time of birth of the young. Another is seen in the early morning rise in adrenal hormones that anticipates wakeup time. Jay's next major review turned to the physiological and evolutionary background of maternal responsiveness (Rosenblatt 1989). Parallel concepts are operative in the analysis of cell autonomous circadian clocks. Thus, a genetic basis of circadian clocks was first identified with the breakthrough work of Konopka and Benzer (1971). They identified three mutants in flies in which the normal 24-hour rhythm was drastically altered. One mutant is arrhythmic; the second had a very short period of about 19 h, and the third had a very long period of 28 h. Each of

these mutations involved the same functional gene on the X chromosome. This was one of the first demonstrations that complex behaviors could be determined by single genes, an observation key to the establishment of the field of behavior genetics. One of Jay's last reviews incorporated contributions of experimental work to the clinical understanding of maternal behavior in humans (Rosenblatt 1994). In work he was contemplating but never finished, Jay was trying to bring these ideas to a more coherent view of evolutionary, genetic and developmental processes. This opportunity to incorporate the findings of basic research and experimentally controlled studies to the problems faced in the clinic was also gaining traction in the field of circadian rhythms. Today, there is tremendous interest in understanding how desynchrony of oscillators in the various tissues of the body underlie illness (Kriegsfeld and Silver 2014). I am sure Jay is delighted with the current interest on epigenetic factors in maternal behavior and in mother–young interactions. He is surely delighted to see the evidence of non-genetic factors tackled at these new levels of analysis. Jay always described the formative effects of experiential factors: It is a thrill to see that they now appear to be accessible to experimental analyses. Rae Silver References Butler, M., Kriegsfeld, L., Silver, R. (2009) Circadian regulation of endocrine functions. In Pfaff, D.W., Arnold, A.P., Etgen, A.M., Fahrbach, S.E., Rubin R.T., eds. Hormones, Brain and Behavior, 2nd edition, Vol 1. San Diego: Academic Press. pp. 473–505. Fleming A.S. (2007) The three faces of Jay S. Rosenblatt. Dev Psychobiol 49: 2–11. Karatsoreos IN, Butler MP, Lesauter J, Silver R. Androgens modulate structure and function of the suprachiasmatic nucleus brain clock. Endocrinology. 2011 May;152(5):1970–8. Konopka RJ and Benzer, S (1971) Clock Mutants of Drosophila melanogaster. PNAS, 68: 2112–2116 Rosenblatt JS. (1980) Hormonal and nonhormonal regulation of maternal behavior: a theoretical survey. Reprod Nutr Dev. 20:791–800. Rosenblatt JS. (1989) The physiological and evolutionary background of maternal responsiveness. New Dir Child Dev. 43: 15–30. Rosenblatt JS, Mayer AD, Giordano AL (1998). Hormonal basis during pregnancy for the onset of maternal behavior in the rat. Psychoneuroendocrinology.13:29–46. Rosenblatt JS. (1994). Psychobiology of maternal behavior: contribution to the clinical understanding of maternal behavior among humans. Acta Paediatr Suppl. 397:3–8. Rosenblatt JS. (2003) Outline of the evolution of behavioral and nonbehavioral patterns of parental care among the vertebrates: critical characteristics of mammalian and avian parental behavior. Scand J Psychol. 44:265–271. Silver R. and Kriegsfeld, L.J. (2014) Circadian Rhythms Have Broad Implications for Understanding Brain and Behavior. 39: 1866–1880. Jay Rosenblatt Jay taught me many things throughout my career but perhaps the most memorable relates to an event that occurred during his undergraduate clinical psychology course. A fellow grad student, Marshall Harth, and I asked to sit in on a few sessions during the semester. During one session, students were presenting their interpretations of the responses to several projective tests that they gave to volunteers. Toward the end of the hour, one student described the responses of a young adult male and then gave a somewhat benign interpretation of the responses. Jay pointed out several disturbing aspects of the male's characterizations of the projective tests. From these, Jay concluded that the male had serious problems forming respectful relationships with women and that he suffered from narcissistic ego-centrism and a lack

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of emotional empathy. The student identified the male as her fiancé and tried to defend him. Jay recommended that she carefully rethink the relationship because it would be very difficult for this male to stay committed emotionally to any female companion. The student asked whether there was anything that could be done to change her fiancé and Jay stated that there was nothing that could be done. Marshall and I confronted Jay after the class. He stated that his interpretation was consistent with the male's responses and we agreed. However, we thought that he was being too pessimistic about the ability of the student to change her fiancé. Couldn't he have offered her some ways of changing the fiancé behavior? Jay replied that there are some aspects of clinical psychology in which we can fully recognize the problem but we do not have any reliable way of “fixing” it. Marshall and I believed that there had to be something that one could do to change such social/emotional characteristics in the fiancé. Subsequently, I learned that Jay was pleased that the couple did not follow his advice and were happily married. Some years later, while I was a member of the psychiatry department at Children's Hospital Medical Center in Boston, I discovered that many brilliant psychiatrists and clinical psychologists were well aware of the weaknesses in the state of the science of changing people and shared Jay's pessimism. However, they were reluctant to communicate such weaknesses to the public (or to granting agencies). Their desire was to never remove hope, even when they could not imagine, much less perceive, how certain social/ emotional problems could be “fixed”. Jay was someone who was very aware of the limits of our knowledge (especially his own). What I learned from him was not to supply a false solution to a problem when a real solution is not available. Although we know too little about the development of psychological characters, we readily use slippery constructs like genetic programs, maturation, neural circuits, learning, culture, etc. to hide our ignorance. Jay's work on maternal behavior was a consistent and programmatic attempt to provide real knowledge to our understanding of how animals (including humans) come to provide care to young. I believe that acknowledging our ignorance is an important quality in a human being and essential for both science and clinical intervention. George F. Michel Zen and the art of maternal cycle maintenance Like the narrator of R.M. Pirsig's (1974) popular and enduring novel, Jay Rosenblatt presented himself with quiet passion, determined inquisitiveness, brutally serious engagement, penetrating perceptions, a delighted esthetic sense, and charisma. Thus, I have borrowed most of the book's title — for fun and with great respect — and use it to recognize and to remember some facets of Jay and his work. Jay was also caring, loving, sensitive and thoughtful. I write as a devotee, someone greatly, nay deeply, affected by Jay and his way. As a contributor of one of the Remembrances, I represent those who never studied formally with Jay and were not students at the Institute for Animal Behavior, but who were nonetheless influenced by Jay as a scientist and teacher. I am grateful to be included in this collection of essays and to share the space with former students and collaborators. As part of a symposium celebrating his 80th birthday, Alison Fleming presented and then published a lovely and illuminating account of the “three faces” of Jay Rosenblatt — scientist, painter, and psychoanalyst (Fleming, 2007). In concert with that broad view of Jay, I had the pleasure of visiting galleries and a museum with Jay, discussing his art, and even learning about bits of his former practice, but here I am remembering him as scientist. Jay's scientific self was big and multifaceted. Appropriately, we are writing these remembrances for the journal, Hormones and Behavior. While Jay certainly advanced knowledge of the hormonal bases of behavior, especially maternal behavior, what I value most from Jay's science is a trove of insights and contributions to a deeper appreciation of

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behavior as a central, organizational topic and for behavioral processes as causal mechanisms. Jay's work was beautiful for its orderliness, logic, and descriptive elegance. His empirical standards matched those of experimental psychology, but he captured and characterized patterns of behavior with rich, multi-leveled analyses more akin to the breadth of ethology. With his friend and colleague, Daniel Lehrman, he provided a lovely and empirically-based description of maternal behavior of the laboratory rat (Rosenblatt & Lehrman, 1963) which, importantly, documented how the behavior changes systematically throughout the postpartum period. This was followed by a rich and important framing of the “synchrony” between the dynamic profile of the mother's behavior and a corresponding suite of changes in the morphology, physiology, sensory function and behavior of the offspring (Rosenblatt, 1965). Rather than seeing behavior in terms of functional units or speciesspecific forms, or with a system of categories, Jay's perspective brought to the fore a view of behavior in time. Maternal behavior is organized as dynamic cycle. When our awareness is expanded to see the organization of behavior as part of continual interactions with the offspring, the concept of synchrony emerges. The term, synchrony, is built on the Greek root, chronos, [time] and this, for me, is emblematic of his commitment to seeing behavior as a phenomenon in time. In this way, it seems to me, Jay also established himself as a developmentalist. He carried his developmental perspective into most of his varied pursuits, including his studies of learning, which inevitably emphasized how learned behaviors develop through time. It is not only the directly observable, ‘superficial’, activities that comprise the material important and relevant to understanding behavior. Jay's probing analyses gave us additional insights into an internal state of readiness to manifest maternal behavior. Here he provided the concept of “maternal responsivity”, admittedly a hypothetical construct, but one that he used masterfully and with his characteristic empirical rigor. Maternal responsiveness was a recurring theme that ran through much of his varied investigations, and those of his many illustrious students. It also became a valuable tool. It is a construct that provides important insight into the organization of behavior. Jay used the quantifiable construct of responsiveness to delineate stages of the maternal behavior cycle — the emergence and dissolution of its expression. He identified phases of the cycle: initiation, maintenance, and decline. Each phase is a kind of entity, defined by the manner in which offspring cues shape it. By experimentally eliminating exposure to offspring stimuli or by exposing mothers to offspring cues of older or younger rat pups, Jay revealed the underlying, active forces that initiate, maintain and decrease the internal state of maternal readiness in the rat dam. Jay and his associates subsequently identified the sensory, endocrine, and experiential bases of each phase and the coordinate transitions between them. This part of his research legacy is thus rich with many important gifts by which he can be justifiably remembered. I probably appreciate most Jay's championing of behavior as the central focus of his analyses, whether his methods seemed to put the work in areas categorized as endocrinological, neural, or learning theory. He was committed to illuminating behavioral processes, and he did it with artistry and high standards of scholarship. Pirsig's novel is often called “an inquiry into values”. Jay's gifts to us are of great value and I think we shall remember them – and him – as a reflection of a life spent inquiring into the ways of giving value to the study of behavior. Jeffrey R. Alberts References Fleming, A. (2007) The three faces of Jay S. Rosenblatt. Developmental Psychobiology, 49(1): 2–11. Pirsig, R.M. (1974) Zen and the Art of Motorcycle Maintenance. New York; William Morrow and Company. Rosenblatt, J.S. (1970) Views on the onset and maintenance of maternal behavior in the rat. In L.R., Aronson, E. Tobach, D.S. Lehrman &

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J.S. Rosenblatt (Eds.), Development and Evolution of Behavior: Essays in Memory of T.C. Schneirla. San Francisco: Freeman, pp. 489–515. Rosenblatt, J.S. (1965) The basis of synchrony in the behavioral interaction between the mother and her offspring in the laboratory rat. In: B.M. Foss (Ed.), Determinants of Infant Behavior, III. New York: John Wiley & Sons. Pp. 3–40. Rosenblatt, J.S. (1967) Nonhormonal basis of maternal behavior in the rat. Science, 156: 1512–1514. Rosenblatt, J.S. & Lehrman, D.S. (1963) Maternal behavior of the laboratory rat. In: H.L. Rheingold (Ed.) Maternal Behavior in Mammals. New York: John Wiley, pp. 8–57. Rosenblatt, J.S., Mayer, A.D., & Siegel, H.I. (1985) Maternal behavior among the non primate mammals. In: Handbook of Behavioral

Neurobiology, vol. 7, N. Adler, D. Pfaff and R.W. Goy (Eds.). New York: Plenum Press, pp. 229–298. Rosenblatt, J.S., Siegel, H.I., & Mayer, A.D. (1979) Progress in the study of maternal behavior in the rat: hormonal nonhormonal, sensory, and developmental aspects. In: J.S. Rosenblatt, R.A. Hinde, C.G. Beer, and M.-C. Busnel (Eds.) Advances in the Study of Behavior, vol. 10. New York: Academic Press. Pp. 226–309.

Jay S. Rosenblatt (November 18, 1923-February 15, 2014).

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