Evolution of passerine incubation behavior: influence of food, temperature, and nest predation

Abstract.— Incubation behavior is one component of reproductive effort and thus influences the evolution of life-history strategies. We examined the relative importance of body mass, frequency of mate feeding, food, nest predation, and ambient temperature to explain interspecific variation in incubation behavior (nest attentiveness, on- and off-bout durations, and nest trips per hour) using comparative analyses for North American passerines in which only females incubate. Body mass and frequency of mate feeding explained little variation in incubation behavior. We were also unable to detect any influence of food; diet and foraging strategy explained little interspecific variation in incubation behavior. However, the typical temperature encountered during reproduction explained significant variation in incubation behavior: Species breeding in colder environments take shorter bouts off the nest, which prevents eggs from cooling to temperatures below the physiological zero temperature. These species must compensate for shorter off-bouts by taking more of them (thus shorter on-bouts) to obtain needed energy for incubation. Nest predation also explains significant variation in incubation behavior among passerines: Species that endure high nest predation have evolved an incubation strategy (long on- and off-bouts) that minimizes activity that could attract predators. Nest substrate explained additional variation in incubation behavior (cavity-nesting birds have shorter on-bouts and make more frequent nest trips), presumably because nest predation and/or temperature varies among nest substrates. Thus, nest predation can influence reproductive effort in a way previously not demonstrated–by placing a constraint on parental activity at the nest. Incubating birds face an ecological cost associated with reproductive effort (predation of entire brood) that should be considered in future attempts to explain avian life-history evolution.     

When to feed: decision making in sticklebacks,Gasterosteus aculeatus

Synopsis The functional analysis of feeding, territory defense and nest-directed activities by reproductively mature male three-spined sticklebacks, Gasterosteus aculeatus, was carried out under controlled laboratory conditions. Territorial males were placed in motivational conflict between feeding and nest-directed activities by making performance of the two necessary activities spatially incompatible. During a limited time each day they had to choose between staying close to their nests to carry out nest-directed activities, and moving into a separate compartment of the observation tank to feed. Males were trapped for varying lengths of time within a central, neutral compartment as they moved between the nest compartment at one end of the tank and the feeding compartment at the other. Their choice of either compartment following such trapping was recorded as indicative of their dominant motivation state. Males tended to be nest dominant in these observations, but motivational dominance could be altered. Males deprived of food for longer periods of time, or presented with an empty food cup during the observation were more likely to be food dominant. If they had not been previously deprived of food, or if a sexually mature conspecific had just been presented in the territory, males were more likely to be nest dominant. The length of time males were trapped in a neutral compartment while in transit between the nest and food also significantly influenced their subsequent behavior. Longer interruptions were more likely to result in a change in the direction in which the male was moving, but only when he was moving towards the subdominant activity. This effect supports the hypothesis of time sharing as the mechanism regulating motivation in these fish.     

Growth rate variation among passerine species in tropical and temperate sites: an antagonistic interaction between parental food provisioning and nest predation risk

Causes of interspecific variation in growth rates within and among geographic regions remain poorly understood. Passerine birds represent an intriguing case because differing theories yield the possibility of an antagonistic interaction between nest predation risk and food delivery rates on evolution of growth rates. We test this possibility among 64 Passerine species studied on three continents, including tropical and north and south temperate latitudes. Growth rates increased strongly with nestling predation rates within, but not between, sites. The importance of nest predation was further emphasized by revealing hidden allometric scaling effects. Nestling predation risk also was associated with reduced total feeding rates and per-nestling feeding rates within each site. Consequently, faster growth rates were associated with decreased per-nestling food delivery rates across species, both within and among regions. These relationships suggest that Passerines can evolve growth strategies in response to predation risk whereby food resources are not the primary limit on growth rate differences among species. In contrast, reaction norms of growth rate relative to brood size suggest that food may limit growth rates within species in temperate, but not tropical, regions. Results here provide new insight into evolution of growth strategies relative to predation risk and food within and among species.     

Spatiotemporal Effects of Supplementary Feeding of Wild Boar (Sus scrofa) on Artificial Ground Nest Depredation

Supplementary feeding of ungulates, being widely used in game management, may have unwanted consequences. Its role in agricultural damage is well-studied, but few studies have considered the potential for the practice to attract ground nest predators. Our goal was to identify the factors influencing ground nest predation in the vicinity of year-round supplementary feeding sites for wild boar and to characterise their spatiotemporal scope. We conducted two separate artificial ground nest experiments in five different hunting districts in south-eastern Estonia. The quantity of food provided and distance of a nest from the feeding site were the most important factors determining predation risk. Larger quantities of food resulted in higher predation risk, while predation risk responded in a non-linear fashion to distance from the feeding site. Although predation risk eventually decreases if supplementary feeding is ceased for at least four years, recently abandoned feeding sites still pose a high predation risk.     

营养改变对潜蝇姬小蜂寄生行为和寄主取食行为的影响

为了研究营养状态对卵育型寄生蜂潜蝇姬小蜂Diglyphus isaea (Walker)雌蜂的寄主取食行为和产卵寄生行为及其二者行为权衡的影响, 在培养皿条件下, 比较了饥饿、加蜂蜜水和不加蜂蜜水3种营养状态的潜蝇姬小蜂雌蜂对美洲斑潜蝇Liriomyza sativae Blanchard各龄幼虫（低、中、高）的寄生、取食及致死能力。结果表明: 非选择条件下, 3种营养状态寄生蜂对高龄幼虫均具有较高的寄生率, 对中龄幼虫具有较高的取食率, 致死率和致死量之间存在显著性差异。3种状态的寄生蜂对低龄幼虫均没有表现出致死能力。有选择条件下, 饥饿状态的寄生蜂对寄主的寄生率最低（5.0%±1.6%）, 取食率最高（16.0%±2.9%）, 特别是对高龄幼虫的取食率占到了整个寄主食物取食率的91.9%; 加蜂蜜水状态下, 寄生蜂对寄主有最低的取食率（8.3%±0.9%）和致死率（17.7%±1.1%）; 不加蜂蜜水状态下, 寄生蜂对寄主有最高的寄生率（13.3%±1.1%）和致死率（28.4%±1.8%）。综合分析发现, 取食寄主的雌蜂比取食蜂蜜水的雌蜂具有更强的致死能力和活动能力。     

Effects of qualitative and quantitative food restriction treatments on feeding motivational state and general activity level of growing broiler breeders

Growing broiler breeder chickens are fed restricted rations to limit body weight at sexual maturity. This experiment tested a proposal (Brouns, F., Edwards, S.A., English, P.R., 1994. Effect of dietary fibre and feeding system on activity and oral behaviour of group housed gilts. Appl. Anim. Behav. Sci. 39, 215-223.) that feeding motivation is reduced by using qualitative rather than quantitative food restriction, and it examined relationships among suppression of growth rate, feeding motivational state and general activity level. From 2 to 15 weeks of age, female broiler breeders were reared in six groups of 20, each with a different feeding treatment. The six treatments were: high and low levels of diet dilution (600-700 g/kg and 300-350 g/kg oat feed, ad libitum), appetite suppression (50-60 g/kg and 25-30 g/kg calcium propionate, ad libitum) and quantitative restriction (recommended daily ration and twice that amount). Birds were conditioned to an operant (PR1 schedule) feeding procedure with their respective treatment diets from 3 to 7 weeks, and this was used to measure feeding motivation in 12-min tests at three times of day (1000, 1300 and 1600 h) at 8, 10, 12 and 14 weeks. Proportions of time spent sitting were measured as an index of general inactivity in systematic observations at 9, 11, 13 and 15 weeks. Although the diet dilution and appetite suppression (qualitative) treatments did not limit growth rates as intended, they and the quantitative treatments produced a range of mean body weights to compare with feeding motivation whenever birds were tested. With body weight as a covariate, there were significant effects on numbers of operant responses in 12 min (the measure of feeding motivation) of weight, age, time of day and treatmentxtime interaction, but not treatment. Feeding motivation was positively correlated with suppression of growth rate, regardless of how that suppression was achieved. However, the experimental procedure required all test birds to be without food from 0900 or 0915 h, and motivation was lowest in the earliest (1000 h) test with qualitative but not quantitative treatments. Hence, there was some evidence that feeding motivation may be partially suppressed with qualitative food restriction; distinction can be made between short-term and longer-term feeding motivation. General activity level (inversely reflected by time spent sitting) was closely correlated positively with both suppression of growth rate and feeding motivation. As a fundamental relationship between feeding motivation and reduction of growth rate was not altered by using qualitative rather than quantitative food restriction, these results support an earlier conclusion (Savory, C.J., Hocking, P.M., Mann, J.S., Maxwell, M.H., 1996. Is broiler breeder welfare improved by using qualitative rather than quantitative food restriction to limit growth rate? Anim. Welfare 5, 105-127.) that broiler breeder welfare is not improved with qualitative restriction methods.     

Compensation for predator-induced reduction in nestling feeding rate in the Great Spotted Woodpecker

Parent birds make efforts to prevent the immediate costs of predation through plastic behavioural responses to the actual predation risk, but this may incur future costs for offspring due to reduced parental care. However, the temporary nature of predator encounters suggests that nestling feeding reduced during the risky periods may be later compensated for by an increased feeding effort (the predation risk allocation hypothesis). We tested this prediction in the Great Spotted Woodpecker (Dendrocopos major) confronted with its major nest predator/competitor, the European Starling (Sturnus vulgaris). A brief encounter with a live starling was followed by a reduction in the nestling feeding rate, but the lost feedings were subsequently compensated for by an increased feeding rate. This compensatory effect was higher in older nestlings that are highly demanding in terms of energy requirements and fitness value from the parents’ perspective. Thus, birds are potentially able to respond not only to the immediate risk of nest predation by nest defence but also to compensate for the potential costs of nest defence in terms of unfulfilled nestling demands. However, data on the amount of delivered food are necessary to distinguish whether parents truly compensate for lost feeding or whether the increased feeding frequency represents a nest guarding strategy.     

Effect of nest and nest site characteristics on the risk of cuckoo Cuculus canorus parasitism in the great reed warbler Acrocephalus arundinaceus

The risk of cuckoo Cuculus canorus parasitism for great reed warbler Acrocephalus arundinaceus nests was evaluated in respect to nest and nest site structure in the reedbeds along canals in central Hungary, In total 90 nests were analysed, from which 31 remained unparasitised. 37 were single parasitised and 22 multiple parasitised. Multiple discriminant analysis, based on eight structural variables, separated well the unparasitised clutches from the cases of single and multiple parasitism, but the latter two categories also showed a weaker separation. Nests close to cuckoo vantage points are most vulnerable to parasitism. The variable "distance from nest to closest available cuckoo perch site" (tree or electric wire) played the most important role in the risk of parasitism. It showed highly significant differences among the groups: lowest distances were found in the case of multiple parasitism, and the highest distances were measured when nests remained unparasitised. Additionally, "nest visibility" also showed significant differences among the three groups: a more visible nest is more likely to be parasitised. Risk of parasitism affects on the host on two levels: 1) female cuckoos search for nest-building hosts from a perch site, but 2) when they are in the act of parasitism, they can find more visible nests in the reed. Besides the robust effects of the variables "distance to cuckoo perch site" and "nest visibility", the variable "distance from nest to open water" may reduce, but "nest volume" increases the risk of multiple parasitism. Differences between cases of single and multiple parasitism are weak, mainly affected by chance. We explain it by the high abundance of the cuckoo, the parasitism rate was 65.6% (59/90), Cuckoos parasitised almost all of the available nests in the close vicinity of potential perch sites. There was no evidence that great reed warblers nested closer to each other when risk of parasitism was high.     

林睡鼠幼鼠的活动规律和行为初步观察

本文通过红外线照相的方法,对发育期的林睡鼠幼鼠进行室内活动规律及行为观察,为充分了解林睡鼠越冬前的能量储备形式提供饲养参考。将鼠密度监测仪固定在饲养笼具上方,24h连续拍照,采集和分析6-9月年龄在10周内的幼鼠各种活动数据。结果显示：幼鼠一天中超过70%的时间都在窝内度过,大部分时间都蜷缩成一团少有动作;在窝外活动时间多在玩耍,如攀爬笼架。用于进食和饮水的时间不超过全天的2%。林睡鼠幼鼠主要活动时间在21:00-07:00,活动高峰在21:00-03:00之间。幼鼠出生3周后开始出窝活动,哺乳期30d后开始采食,5周后幼鼠有交配玩耍行为。随着日龄的增长,活动高峰从23：00提前到21：00,活动时间也逐渐延长,但9周龄后活动时间逐渐缩短,幼鼠的饮水进食时间与其活动的时间长短较为一致。10周体重可达成年体重的70%。研究表明,林睡鼠在夏秋季节基本是昼伏夜出动物。光照是影响其在外活动的重要因素之一。幼鼠6周后所需的饲料和水量大于成年林睡鼠,在此期间要注意饲料和水的补充。     

Feeding ecology of the earliest vertebrates

Based on protochordates and extant fish, the earliest Palaeozoic vertebrates were microphagous suspension-feeding animals that pumped food-carrying water very slowly and thus required highly concentrated suspensions. Such conditions exist in benthic (not open water) aquatic environments. Feeding modes which on the basis of extant fish are closely related to benthic microphagous suspension feeding include deposit feeding, epilithic algal scraping, and macrophagous suspension feeding; early jawless vertebrates are predicted to have included all these feeding types. The gnathostome condition is predicted to have followed an initial switch from feeding on suspensions to taking tiny individual food particles (microphagous suspension-feeding → microphagous particulate-feeding → macrophagous particulate-feeding).     

Gastrin in the mechanisms of genetic determination of feeding behavior in rabbits

Spreading of a dominant motivation to the molecular intracellular mechanisms of genetic memory was studied. Blockage of protein synthesis in the nervous system selectively abolishes food motivation in rabbits during stimulation of the lateral hypothalamus but exerts no noticeable effect on avoidance responses during stimulation of the ventromedial hypothalamus. During protein synthesis blockage, food motivation returns to normal upon pentagastrin intracerebroventricular (i.c.v.) injection. Intracerebroventricular administration of antigastrin immunoglobulin inhibits feeding reaction to lateral hypothalamic stimulation but not avoidance response to ventromedial hypothalamic stimulation. It was concluded that feeding motivation translates into feeding behavior in the following stages: motivational excitation, gene activation, mRNA synthesis, formation of a gastrin-like peptide, and expression of feeding behavior.     

Temperature and forager body size affect carbohydrate collection in German yellowjackets, Vespula germanica (Hymenoptera, Vespidae)

The classic formulation of optimal foraging theory predicts that a central-place forager will gather more food if it is required to travel farther from the nest to find that food. We examined the foraging behavior of German yellowjackets (Vespula germanica) to determine whether carbohydrate foragers follow this pattern. We trained foragers to collect 2 M fructose solution at 5 or 50 m from the nest and measured the time spent feeding, load size, and the rate of delivery. We show that as a forager’s crop fills during a foraging bout, the amount of solution ingested per second decreased. However, load size did not change as wasps collected food up to 50 m from the nest. Instead, temperature and body size were better predictors of the volume of fructose a forager carried. Finally, the rate of fructose delivered to the nest was higher at warmer temperatures. Due to the fact that wasps gather more food but feed for shorter periods of time at warmer temperatures, we found an overall negative relationship between feeding time and load size. We conclude that the strong effects temperature had on the behavior of V. germanica foragers imply that feeding time may not always be an accurate predictor of the size of the load an individual carries back to the nest. Results from this study suggest that in yellowjacket colonies, foragers can collect and bring disproportionately more food back to the nest during the warmest days of the summer, a time of year when this pest species reaches peak population size during its annual colony cycle.     

Incubation feeding by male Scarlet Tanagers: a mate removal experiment

ABSTRACT Incubation feeding, where males feed their mates, is a common behavior in birds and may improve female condition, nest attentiveness, and nesting success. We used behavioral observations and a temporary mate removal experiment to test the female nutrition hypothesis for incubation feeding by male Scarlet Tanagers (Piranga olivacea). All males (N= 20) were observed incubation feeding and fed females both at the nest (x? 1.36 trips/h) and away from the nest (x? 20.1 trips/h). Male feeding rate off‐nest was negatively correlated with the duration of female foraging bouts and positively correlated with the total time females spent incubating per hour. Eggs were predated at seven of 19 (37%) nests, but nest survival during incubation was not related to either female incubation behavior or male feeding rate. During temporary removal experiments (N= 12), female Scarlet Tanagers remained on the nest significantly longer and did not have longer foraging bouts. An unexpected outcome of the removal experiments was a dramatic change in female vocal behavior. All 12 experimental females gave chik‐burr calls during the male‐removal experiments (x? bout length = 11.7 min), but during normal observation periods only six of 20 females at the incubation stage gave chik‐burr calls (x? bout length = 0.7 min, N= 20). Our results suggest that female tanagers likely gain nutritional benefits from incubation feeding, but male feeding may not improve immediate reproductive success. Nine of 54 (17%) nestlings in five of 17 broods (29%) were extra‐pair young (EPY), indicating that males could potentially benefit from incubation feeding via mate retention and fidelity as well as, or instead of, through immediate gains in reproductive success. Our study indicates that females benefit from incubation feeding and do not simply passively accept food from their mates, but instead may influence male feeding rates through direct (e.g., mate following and vocalizing) and indirect (the threat of mate abandonment or cuckoldry) means.     

Regulation of feeding behavior and food intake by appetite-regulating peptides in wild-type and growth hormone-transgenic coho salmon

Survival, competition, growth and reproductive success in fishes are highly dependent on food intake, food availability and feeding behavior and are all influenced by a complex set of metabolic and neuroendocrine mechanisms. Overexpression of growth hormone (GH) in transgenic fish can result in greatly enhanced growth rates, feed conversion, feeding motivation and food intake. The objectives of this study were to compare seasonal feeding behavior of non-transgenic wild-type (NT) and GH-transgenic (T) coho salmon (Oncorhynchus kisutch), and to examine the effects of intraperitoneal injections of the appetite-regulating peptides cholecystokinin (CCK-8), bombesin (BBS), glucagon-like peptide-1 (GLP-1), and alpha-melanocyte-stimulating hormone (α-MSH) on feeding behavior. T salmon fed consistently across all seasons, whereas NT dramatically reduced their food intake in winter, indicating the seasonal regulation of appetite can be altered by overexpression of GH in T fish. Intraperitoneal injections of CCK-8 and BBS caused a significant and rapid decrease in food intake for both genotypes. Treatment with either GLP-1 or α-MSH resulted in a significant suppression of food intake for NT but had no effect in T coho salmon. The differential response of T and NT fish to α-MSH is consistent with the melanocortin-4 receptor system being a significant pathway by which GH acts to stimulate appetite. Taken together, these results suggest that chronically increased levels of GH alter feeding regulatory pathways to different extents for individual peptides, and that altered feeding behavior in transgenic coho salmon may arise, in part, from changes in sensitivity to peripheral appetite-regulating signals.     

Food division in the Arabian babbler nest: adult choice or nestling competition?

Many studies have found that food division among nestmates isnegatively correlated with hatrhing order. The hypothesis thatfeedings are distributed according to the provisioners' selectionis compared here with the hypothesis that feedings are dividedas a consequence of nestling competition. Food distributionamong Arabian babbler nestmates was negatively correlated withhatching order and was so skewed that many last-hatched nestlingsdied as a consequence of starvation. No feeding preference wasfound between any adult and any given nestling. No differencewas found in feeding distribution among parents, partial parents,and nonparents. When an older foreign nestling was introducedinto the nest, adults fed the newcomer more intensively thantheir own offspring, implying that feeding division among nestmateswas not determined by die feeders but by the nestlings' relativeability to obtain food. In two-nestling nests, the older nestlingobtained 62% of the feedings, on average. However, followingselective feedings of the second-hatched nestling by the investigator,which equalized the two siblings' feeding amounts by the endof their second day, no difference was found between their feedingrates. The relative advantage of the older nestlings was eliminatedby introducing a barrier into the nest. Under these conditions,feedings were distributed equally among the nestmates. Sinceneither hatching order, previous priority order, nor individualidentification were used by the provisioning adults, the proximatefactor determining food division among the neslings appearsto be the result of nestling competition     

Predator-induced plasticity in nest visitation rates in the Siberian jay (Perisoreus infaustus)

Bird nestlings may be at risk not only from starvation but alsofrom predators attracted to the nest by parental feeding visits.Hence, parents could trade reduced visitation rates for a lowerpredation risk. Here, through field data and an experiment,we show plasticity in daily patterns of nest visitation in theSiberian jay, Perisoreus infaustus, in response to predatoractivity. In high-risk territories, jay parents avoided goingto the nest at certain times of the day and compensated by allocatingmore feeding effort to periods when predators were less active.Such modifications in provisioning routines allowed parentsin high-risk habitat to significantly lower the risk of providingvisitation cues to visually oriented corvid nest predators.These results indicate that some birds modify their daily nestvisitation patterns as a fourth mechanism to reduce predator-attractingnest visits in addition to the clutch size reduction, maximizationof food load-sizes, and prevention of allofeeding suggestedby Skutch.     

Nest site and weather affect the personality of harvester ant colonies

Environmental conditions and physical constraints both influence an animal's behavior. We investigate whether behavioral variation among colonies of the black harvester ant, Messor andrei, remains consistent across foraging and disturbance situations and ask whether consistent colony behavior is affected by nest site and weather. We examined variation among colonies in responsiveness to food baits and to disturbance, measured as a change in numbers of active ants, and in the speed with which colonies retrieved food and removed debris. Colonies differed consistently, across foraging and disturbance situations, in both responsiveness and speed. Increased activity in response to food was associated with a smaller decrease in response to alarm. Speed of retrieving food was correlated with speed of removing debris. In all colonies, speed was greater in dry conditions, reducing the amount of time ants spent outside the nest. While a colony occupied a certain nest site, its responsiveness was consistent in both foraging and disturbance situations, suggesting that nest structure influences colony personality.     

杂色山雀亲代喂食与子代乞食间的行为

在育雏期,晚成鸟的子代一般都是由双亲共同来抚育,子代为了更好地存活,会用自己的方式竞争获得更多的食物和更好的生存空间,同时亲代也会根据子代的乞食信号来分配食物。2011年3～7月采用针孔摄像技术录制了杂色山雀(Parus varius)育雏期巢内亲代与子代间的行为,统计了亲鸟站位、雏鸟站位、雏鸟乞食强度及亲鸟的喂食情况等数据。分析结果表明:(1)雌雄亲鸟在巢中的站位各有特点,雄鸟在整个育雏期都喜欢站在距离巢口较近的位置;雌鸟站位不太固定,前期离巢口相对较远,中期和后期离巢口相对较近;(2)雏鸟离亲鸟越近,乞食强度越大,获得食物的机会就越多;离亲鸟越远的雏鸟越不爱乞食,所以站位对雏鸟的食物获得影响最大;(3)雌鸟承担主要的育雏任务,喂食频率远大于雄鸟;(4)育雏期的不同阶段雏鸟乞食强度、亲鸟喂食频率变化很大:中期雏鸟乞食强度最大,亲鸟喂食频率最高,后期雏鸟乞食强度最弱;(5)整个育雏期雌性亲本没有表现出明显的偏爱行为,但雄性亲本在中、后期更偏爱体型大的雏鸟。可见杂色山雀子代的行为和体型大小影响着亲代的食物分配,亲代也会根据雏鸟日龄调整站位和喂食行为。     

Predatory behavior of a seed-eating ant:Brachyponera senaarensis

The great flexibility of the feeding strategies exhibited by the ponerine ant Brachyponera senaarensis (Mayr) allows it to exploit either seeds or animal prey items as food resources. Predation is generally limited to small prey and is very similar to scavenging behavior. In laboratory conditions, the predatory behavior of B. senaarensis is not different in structure from that known in other carnivorous ants species. The workers forage individually and return to the nest using a series of cues involving light, a chemical graduated marking system near the nest entrance, and memory. During nest-moving, recruitment by tandem running was observed. However, in colonies where the food supply is regular, workers that discover food do not recruit nestmates, but make repeated trips between the nest and the food source. On the contrary, in starved colonies, the introduction of prey may produce a massive exit of foragers, corresponding to a primitive form of mass recruitment similar to that observed in some other ant species.     

Identification of neurons responsible for feeding behavior in the Drosophila brain

Drosophila melanogaster feeds mainly on rotten fruits,which contain many kinds of sugar.Thus,the sense of sweet taste has evolved to serve as a dominant regulator and driver of feeding behavior.Although several sugar receptors have been described,it remains poorly understood how the sensory input is transformed into an appetitive behavior.Here,we used a neural silencing approach to screen brain circuits,and identified neurons labeled by three Gal4 lines that modulate Drosophila feeding behavior.These three Gal4 lines labeled neurons mainly in the suboesophageal ganglia(SOG),which is considered to be the fly’s primary taste center.When we blocked the activity of these neurons,flies decreased their sugar consumption significantly.In contrast,activation of these neurons resulted in enhanced feeding behavior and increased food consumption not only towards sugar,but to an array of food sources.Moreover,upon neuronal activation,the flies demonstrated feeding behavior even in the absence of food,which suggests that neuronal activation can replace food as a stimulus for feeding behavior.These findings indicate that these Gal4-labeled neurons,which function downstream of sensory neurons and regulate feeding behavior towards different food sources is necessary in Drosophila feeding control.