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Fitness consequences of hoarding behaviour in the Eurasian red squirrel.

Hoarding increases food availability during periods of scarcity, and therefore should enhance fitness. Although short-term advantages of hoarding have been described for birds, effects over an animal's lifetime have not yet been documented. Here, we report that in the red squirrel, Sciurus vulgaris, individuals which recovered many cached tree seeds increased their body mass and were more likely to survive the spring breeding season than those that recovered fewer seeds. There was no significant effect of the time spent recovering cached food on the probability for females to produce a spring litter. In the long-term, hoarding behaviour was related to fitness in two ways; (i) squirrels spending more time recovering hoards survived longer; and (ii) females with a high recovery activity tended to wean more young in their lifetime than those that spent less time recovering hoards. Our data indicate that in red squirrels, food hoarding is an adaptive foraging strategy to preserve temporarily abundant food resources for future periods of hardship, and that individuals that hoard and recovery many tree seeds are more likely to survive and reproduce.

Adaptation, Physiological↗

Is hippocampal volume affected by specialization for food hoarding in birds?

The hypothesis that spatial-memory specialization affects the size of the hippocampus has become widely accepted among scientists. The hypothesis comes from studies on birds primarily in two families, the Paridae (tits, titmice and chickadees) and the Corvidae (crows, nutcrackers, jays, etc.). Many species in these families store food and rely on spatial memory to relocate the cached items. The hippocampus is a brain structure that is thought to be important for memory. Several studies report that hoarding species in these families possess larger hippocampi than non-hoarding relatives, and that species classified as large-scale hoarders have larger hippocampi than less specialized hoarders. We have investigated the largest dataset on hippocampus size and food-hoarding behaviour in these families so far but did not find a significant correlation between food-hoarding specialization and hippocampal volume. The occurrence of such an effect in earlier studies may depend on differences in the estimation of hippocampal volumes or difficulties in categorizing the degree of specialization for hoarding or both. To control for discrepancies in measurement methods we made our own estimates of hippocampal volumes in 16 individuals of four species that have been included in previous studies. Our estimates agreed closely with previous ones, suggesting that measurement methods are sufficiently consistent. Instead, the main reasons that previous studies have found an effect where we did not are difficulties in assessing the degree of hoarding specialization and the fact that smaller subsets of species were compared than in our study. Our results show that a correlation between food-hoarding specialization and hippocampal volume cannot be claimed on the basis of present data in these families.

Adaptation, Physiological↗

Agouti-related protein increases food hoarding more than food intake in Siberian hamsters.

Agouti-related protein (AgRP), an endogenous melanocortin 3/4 receptor antagonist, appears to play an important role in the control of food intake and energy balance because exogenous administration in rats and overexpression in mice result in hyperphagia and body mass gain. Furthermore, arcuate nucleus AgRP mRNA is increased with fasting in laboratory rats and mice and is decreased with refeeding. In Siberian hamsters, fasting also increases arcuate nucleus AgRP mRNA, but these animals increase food hoarding, rather than food intake with refeeding. Therefore, we tested whether exogenous AgRP increased food hoarding in this species. Hamsters were trained in a hoarding/foraging apparatus to run a programmed number of wheel revolutions to earn food pellets. Four doses of AgRP-(83-132) or vehicle were injected into the third ventricle at the beginning of the dark phase, and food hoarding, food intake, and foraging were measured at various time points subsequently. Overall, food hoarding was stimulated as much as 10 times more than food intake, and both responses occurred as early as 1 h after injection. Food hoarding was increased the greatest at the lowest dose (0.1 nmol), whereas food intake was increased the greatest at the second lowest dose (1 nmol). Food intake and especially food hoarding were increased up to seven days after the AgRP injections. Foraging was increased at all AgRP doses except the highest dose (100 nmol). These results suggest that AgRP triggers the search for food in this species, and once they find it, hoarding predominates over eating.

Agouti-Related Protein↗

NPY Y1 receptor is involved in ghrelin- and fasting-induced increases in foraging, food hoarding, and food intake.

Fasting triggers a constellation of physiological and behavioral changes, including increases in peripherally produced ghrelin and centrally produced hypothalamic neuropeptide Y (NPY). Refeeding stimulates food intake in most species; however, hamsters primarily increase foraging and food hoarding with smaller increases in food intake. Fasting-induced increases in foraging and food hoarding in Siberian hamsters are mimicked by peripheral ghrelin, central NPY, and NPY Y1 receptor agonist injections. Because fasting stimulates ghrelin and subsequently NPY synthesis/release, it may be that fasting-induced increased hoarding is mediated by NPY Y1 receptor activation. Therefore, we asked: Can an Y1 receptor antagonist block fasting- or ghrelin-induced increases in foraging, food hoarding, and food intake? This was accomplished by injecting the NPY Y1 receptor antagonist 1229U91 intracerebroventricularly in hamsters fasted, fed, or given peripheral ghrelin injections and housed in a running wheel-based food delivery foraging system coupled with simulated-burrow housing. Three foraging conditions were used: 1) no running wheel access, free food, 2) running wheel access, free food, or 3) foraging requirement (10 revolutions/pellet) for food. Fasting was a more potent stimulator of foraging and food hoarding than ghrelin. Concurrent injections of 1229U91 completely blocked fasting- and ghrelin-induced increased foraging and food intake and attenuated, but did not always completely block, fasting- and ghrelin-induced increases in food hoarding. Collectively, these data suggest that the NPY Y1 receptor is important for the effects of ghrelin- and fasting-induced increases in foraging and food intake, but other NPY receptors and/or other neurochemical systems are involved in increases in food hoarding.

Animals↗

Rats eating and hoarding as a function of body weight and cost of foraging.

Rats were trained to forage 2 h daily in a zigzag alley at -15 degrees C. On experimental days food was placed in large amounts at a 1-, 4-, or 16-m distance from a warm shelter. The mass of food ingested and hoarded in the shelter by the rats was then recorded. The rats' body weights were depressed progressively by spacing the sessions with access to food, and then the animals were allowed to recover their initial body weight. The results showed that 2-h food intake was constant and did not vary as a function of body weight loss or the distance to reach the food (cost). On the other hand, the mass of food hoarded was a linear increasing function of body weight loss below the hypothetical set point for body weight and a decreasing linear function of the distance to the food. The linear regression of the mass of food hoarded on body weight with a negative slope (b) adequately depicted the rat's hoarding behavior: mass hoarded = a + b (body wt). The slope of this function decreased with increasing cost of food procurement. It is concluded that 1) the main response of the rat to starvation is food hoarding rather than ingestion and 2) the estimation of the body weight set point from hoarding is not affected by the costs of food procurement.

Animals↗

Partial lipectomy, but not PVN lesions, increases food hoarding by Siberian hamsters.

We tested the inverse relationship between body fat and food hoarding in Siberian hamsters by decreasing or increasing body fat through partial surgical lipectomy (LIPX) or by making obesity-inducing lesions of the paraventricular nucleus of the hypothalamus (PVNx), respectively. We asked three questions. 1) Is food hoarding increased after body fat loss due to LIPX? 2) Is food hoarding decreased after PVNx? 3) Does PVNx affect the hoarding response to LIPX? Hamsters housed in a simulated burrow system increased food hoarding after LIPX followed by a decrease to pre-LIPX levels as body fat was partially compensated through an increase in the mass of their unoperated fat pads. PVNx hamsters had increased body mass and food intake but did not have decreased food hoarding, nor was food hoarding increased by LIPX in PVNx hamsters. The partial body fat compensation by LIPX + PVNx hamsters suggests that the damaged PVN did not cause a general failure to sense energy deficits but did affect the ability to integrate internal and external energy stores.

Adipose Tissue↗

Cerebral glucose metabolism in obsessive-compulsive hoarding.

OBJECTIVE: Compulsive hoarding and saving symptoms, found in many patients with obsessive-compulsive disorder (OCD), are part of a discrete clinical syndrome that includes indecisiveness, disorganization, perfectionism, procrastination, and avoidance and has been associated with poor response to medications and cognitive behavior therapy. The authors sought to identify cerebral metabolic patterns specifically associated with the compulsive hoarding syndrome using positron emission tomography (PET). METHOD: [(18)F]Fluorodeoxyglucose PET scans were obtained for 45 adult subjects who met DSM-IV criteria for OCD (12 of whom had compulsive hoarding as their most prominent OCD symptom factor) and 17 normal comparison subjects. All subjects had been free of psychotropic medication for at least 4 weeks. Regional cerebral glucose metabolism was compared between the groups. RESULTS: In relation to the comparison subjects, the patients with compulsive hoarding syndrome had significantly lower glucose metabolism in the posterior cingulate gyrus and cuneus, whereas the nonhoarding OCD patients had significantly higher glucose metabolism in the bilateral thalamus and caudate. In relation to nonhoarding OCD patients, compulsive hoarders had significantly lower metabolism in the dorsal anterior cingulate gyrus. Across all OCD patients, hoarding severity was negatively correlated with glucose metabolism in the dorsal anterior cingulate gyrus. CONCLUSIONS: OCD patients with the compulsive hoarding syndrome had a different pattern of cerebral glucose metabolism than nonhoarding OCD patients and comparison subjects. Obsessive-compulsive hoarding may be a neurobiologically distinct subgroup or variant of OCD whose symptoms and poor response to anti-obsessional treatment are mediated by lower activity in the cingulate cortex.

Adult↗

Hoarding and obsessive-compulsive symptoms.

The present study attempts to extend recent research on the relation between hoarding and obsessive-compulsive experiences. In both college student and community samples, hoarding was associated with higher scores on the Yale-Brown Obsessive-Compulsive Scale (YBOCS). The relationship was stronger among the community sample, in which there was a greater range of compulsive symptoms and hoarding behavior. Hoarding was also associated with higher levels of general psychopathology as measured by the Brief Symptom Inventory but not by the Obsessive-Compulsive Personality Disorder subscale from the Millon Clinical Multiaxial Inventory-II or by a measure of ordinary risk taking. Among a sample of patients with obsessive-compulsive disorder (OCD), 31% reported hoarding obsessions and 26% reported hoarding compulsions on the YBOCS symptom checklist. These frequencies are similar to those found elsewhere and suggest that, although not as frequent as the classical symptoms of OCD, hoarding is a common symptom among OCD patients.

Adolescent↗

Daily hoarding opportunity entrains the pacemaker for hamster activity rhythms.

The effects on activity rhythms of a daily 30 min opportunity to leave the home cage and hoard seeds from an open field were assessed in Syrian hamsters housed in continuous dim illumination. Six of ten hamsters responded with clear entrainment of their activity rhythms to the hoarding opportunity, as demonstrated by responses to phase shifts and by the onset phase of subsequent freerunning rhythms. No entrainable component separate from the freerunning rhythm was ever observed. Two hamsters showed phase shifts in response to the hoarding opportunity, but they did not meet the criteria for stable entrainment, and two did not respond with noticeable changes in rhythmicity. Ablations of the suprachiasmatic nuclei (SCN) were attempted in three hamsters that had entrained stably to the hoarding time. The effects of partial lesions in two animals indicated that the entrained rhythm was controlled by the light-entrainable pacemaker represented by the SCN. The one animal with an apparently complete lesion, however, developed a clear, but irregular, increase in activity in anticipation of the daily hoarding time. SCN ablation apparently unmasked an oscillator system separate from the SCN and susceptible to entrainment by a nonphotic cue. The oscillator mechanism affected by daily hoarding opportunities in hamsters appears to be tightly coupled to the SCN pacemaker, in contrast to the system in rats that is synchronized by daily feeding schedules.

Animals↗

Hoarding behavior of AKR, C57BL/6 mice and their F1 in their home living space: an automatic recording technique.

Earlier genetic analysis of food hoarding behavior in mice used test situations in which animals were allowed to hoard food only during short time periods. This paper describes a technique to measure food hoarding over long time periods. Small discs of standard food are equipped with a steel clip and each passage of a mouse bringing such a food disc from the food search enclosure to the nest is detected by an electromagnetic sounding lead connected to a pen recorder. Preliminary results show that female mice begin to hoard earlier than males; one of the two reciprocal F1 has a shorter latency to hoard than the other one. Results also show differences between two parental inbred strains and their reciprocal F1 for the light-dark distribution of hoarding.

Animals↗

Effect of a cold ambient temperature on the rat's food hoarding behavior.

In a first experiment, the amount of food hoarded in three hour sessions (HB) by eight food deprived rats, was measured at three ambient temperatures: 30, 17.5 and 5 degrees C. At the three temperatures, the mass of food pellets hoarded by the rats was linearly related to the rats' body weight (BW) with a negative slope. The slope of the regression lines was significantly higher at 5 and 17.5 degrees C than at 30 degrees C. In a second experiment, BW and HB of ten non-deprived rats were measured every day, first at 30, then at 10 degrees C, then at 30 degrees C again (70 days). In the cold environment, HB immediately increased without BW loss. This effect was immediately reversible by the adjunction of an insulating styrofoam mattress. HB of non deprived rats in the cold was therefore more a thermoregulatory behavior than an alimentary response. In a third experiment, ad lib fed or deprived rats could hoard either food pellets or styrofoam lumps at 30 degrees C or at 5 degrees C. In the cold environment, when non-deprived, the rats hoarded styrofoam rather than food. However, as soon as they were food deprived, they reversed their choice and hoarded food rather than styrofoam. It is concluded that food can be hoarded as a thermal insulator in a cold environment but HB is mainly an alimentary response increased by a cold stress.

Animals↗

Acute intraventricular CRF lowers the hoarding threshold in male rats.

The influence of intracerebroventricular rat-CRF on the food-hoarding behavior of rats has been studied in relation to the animals' body weights. A group of six male rats was trained to feed every day from 1000 to 1200 h. Then their threshold for the onset of food hoarding was measured from the intercept of regression line of food hoarded during meal time vs. body weight with the x-axis. Thirty minutes before the hoarding session, the rats received 4 micrograms CRF, or saline control, in the lateral ventricle. The mean threshold for food hoarding was significantly lowered to 299 +/- 61 g after CRF, from control 418 +/- 68 g. Mean food intake during the hoarding sessions was also diminished to 6.0 +/- 0.6 g after CRF, from control 14.5 +/- 0.5 g. These results suggest that the set-point for body weight regulation is acutely lowered by intracerebral CRF.

Animals↗

Food hoarding and ingestion in the deer mouse, Peromyscus maniculatus: selective responses to mu and kappa opiate agonists.

The feeding behavior of the deer mouse, Peromyscus maniculatus, includes food hoarding as well as ingestion. Administration of the prototypical mu opiate agonist, morphine sulfate, 1-20 mg/kg, produced over three hours a significant dose-dependent stimulation of hoarding by free feeding deer mice. The specific kappa opiate agonist, U-50,488H, 0.10-10 mg/kg, markedly increased ingestion without having any augmentatory effects on hoarding. The mixed mu and kappa opiate agonist, ketocyclazocine hydrochloride, 1-10 mg/kg, as well as various combinations of morphine sulfate and U-50,488H, augmented both hoarding and ingestion. Food restriction for 24 hr caused a significant, naloxone (1.0 mg/kg) reversible, increase in food intake. Food deprivation also modified the hoarding and ingestion responses of the deer mice to the mu and kappa opiate agonists, reducing the relative amounts of food that were hoarded. These results indicate that mu and kappa opioid systems are differentially involved in the mediation of various aspects of feeding. This also suggests that environmental factors, such as food restriction, can modify the relative roles of mu and kappa opioid systems in the expression of feeding behavior.

3,4-Dichloro-N-methyl-N-(2-(1-pyrrolidinyl)-cycloh↗

The hoarding dimension of OCD: psychological comorbidity and the five-factor personality model.

Although hoarding has been associated with several psychological disorders, it is most frequently linked to obsessive-compulsive disorder (OCD). The present study assessed hoarding obsessions and compulsions in 204 individuals with OCD, and evaluated how hoarding was related to obsessive-compulsive symptom severity, psychological comorbidity, and personality as measured by the five-factor model. Results indicated that hoarding in OCD is a dimensional variable that is positively associated with dysphoria, total number of lifetime Axis I disorders, and lifetime histories of bipolar I, PTSD, and body dysmorphic disorder. Hoarding was negatively correlated with the NEO-Personality Inventory-Revised (NEO-PI-R) factor of Conscientiousness and positively associated with the NEO-PI-R factor of Neuroticism. When all personality and psychopathology variables were entered into a regression equation, dysphoria, bipolar II disorder, Conscientiousness, age, and Extraversion emerged as significant predictors of hoarding severity. Recommendations are made for clinicians and for future research.

Adolescent↗

Diet choice exaggerates food hoarding, intake and pup survival across reproduction.

Siberian hamsters increase food intake and hoarding during pregnancy and lactation, perhaps to compensate for large decreases in body fat (approximately 50%). We tested the effects of diet choice on these responses in pregnant, lactating and virgin hamsters housed in a simulated burrow system. Hamsters were offered pellet chow (PC) or a choice of sunflower seeds (SS), rabbit chow (RC) and PC. Pregnant or lactating PC-fed hamsters had increased food intake and hoard size compared with virgins, effects exaggerated by diet self-selection. The pregnancy-induced increases and lactation-induced decreases in body mass were enhanced and diminished by diet self-selection, respectively. Pregnant self-selecting hamsters ate relatively more carbohydrate and less fat and hoarded less carbohydrate and more fat than their virgin counterparts (protein not affected). Lactating and virgin self-selecting hamsters both ate and hoarded relatively more carbohydrate than protein or fat compared with PC-fed hamsters but were not different from each other. Litter and pup sizes were similar at birth, but pups from self-selecting mothers had decreased lipid content (50%) compared with pups from PC-fed mothers, whereas at weaning they were heavier but not fatter. Only lactating PC-fed mothers cannibalized their pups (approximately 60% eaten, 8/10 litters). The pregnancy-induced increased eating and hoarding of carbohydrate may have helped meet immediate energy needs sparing dwindling lipid reserves, whereas the decreased fetal lipid investment may have helped conserve energy in anticipation of the increased demands of lactation. The diet-induced exaggerated caloric intake and food hoard size of lactating hamsters may have promoted pup growth and survival.

Adipose Tissue↗

The food-hoarding threshold is not raised by acute intraventricular NPY in male rats.

We studied the influence of acute intracerebroventricular (i.c.v.) NPY on the relation existing in the rat between food-hoarding behavior and the body weight of the animals. Six male rats were trained to feed every day from 1000 to 1200 h. Then, their threshold for the onset of food hoarding was measured from the intercept of the regression line relating the food hoarded during meal time (Y axis) to the body weight (X axis). Thirty minutes before the hoarding session, the rats were injected i.c.v. with either 4 micrograms of NPY or saline control. The mean threshold for food hoarding was not modified after NPY. Mean food intake during the hoarding sessions was also unchanged. These results suggest that intracerebral NPY at 4 micrograms did not acutely alter the set-point for body weight regulation.

Animals↗

Disappearance of hoarding behavior after 6-hydroxydopamine lesions of the mesolimbic dopamine neurons and its reinstatement with L-dopa.

The consequences of 6-hydroxydopamine lesions of the mesolimbic dopamine system on hoarding behavior were investigated in the rat. Specific lesions of this system, at the level of either the ventral tegmental area or the nucleus accumbens, resulted in abolition or severe reduction of hoarding activity. Similar lesions of the forebrain noradrenaline neurons did not affect hoarding. In further experiments, amphetamine and apomorphine locomotor responses, spontaneous motor behavior, food intake and eating patterns, and the existence of any regulatory deficits were examined. A subtle disorganization of eating patterns was found in animals with mesolimbic-dopamine lesions. It was determined that the hoarding deficit could not be due to motor, ingestive, or regulatory impairments. In a final experiment, it was demonstrated that hoarding behavior can be restored to control levels in dopamine-lesion rats by prior treatment with the catecholamine presursor L-dopa. These findings suggest that hoarding activity is mediated by mesolimbic dopamine neurons, and it is hypothesized that this system is necessary for the facilitation of certain types of foraging responses under a high level of arousal.

Animals↗

On phenomenology and classification of hoarding: a review.

OBJECTIVE: Hoarding is a behavioural abnormity characterized by the excessive collection of poorly usable objects. It is described mainly in association with obsessive-compulsive disorders (OCDs) and in geriatric populations. Yet the literature on the phenomenon is heterogeneous and the notion obviously lacks a consistent definition. This review attempts to describe the psychopathological and clinical spectrum of hoarding and may contribute to clarify its classification. METHOD: Systematic review and discussion of the literature on hoarding. RESULTS: Hoarding is a complex behavioural phenomenon associated with different mental disorders. The psychopathological structure is variously composed of elements of OCDs, impulse-control disorders, and ritualistic behaviour. Severe self-neglect is a possible consequence of hoarding. CONCLUSION: Without further specifications the term hoarding is of limited heuristic value and cannot guide therapeutic interventions satisfactorily. The condition needs to be evaluated carefully in every particular case in relation to the aforementioned psychopathological concepts.

Aged↗