Diurnal variation in heat production related to some physical activities in laying hens

1. Heat production, standing and eating activities, and hourly food intake of 4 laying hens were observed simultaneously and the effects of activity and food intake on heat production were studied. 2. Average heat production during the dark period (20.00 to 06.00 h) was 18.9 kJ/kgW0.75 h which was 33% lower than that during the light period. About 76% of the light-dark difference in the rate of heat production was probably associated with activity and posture. 3. Standing time, which included a range of behavioural activities, occupied 90% of the light period and the increased rate of heat production associated with standing was estimated to be about 18% of daily heat production. 4. Eating time occupied 40% of the light period; the heat production associated with eating activity represented about 5% of daily heat production or 3% of ME intake. 5. Because the regression of heat production on time spent eating agreed with the regression of heat production on hourly food intake, it is suggested that the energy expenditure associated with ad libitum feeding can be estimated for hens from the regression of heat production on hourly food intake.     

Effects of environmental temperature on heat production associated with food intake and on abdominal temperature in laying hens.

An experiment was carried out to verify the relationship between the heat production associated with food intake and environmental temperature. Four laying hens were trained to eat two meals per day and were kept under artificial illumination with dark, dim and light periods. 2. Metabolic heat production was measured using two open-circuit respiratory chambers. Abdominal temperatures were measured, using thermocouples, at environmental temperatures of 12, 16, 20, 24, 28, 32 and 36 degrees C and at food intakes of 90, 60, 30 and 0 g/d. 3. The rate of heat production was dependent on both environmental temperature and food intake. Increasing environmental temperature resulted in a decrease in total metabolic rate at any food intake, indicating that heat production associated with food intake was not directly linked with thermoregulation at low environmental temperatures. 4. Abdominal temperature varied little with either food intake or environmental temperature below 28 degrees C. Above 28 degrees C, abdominal temperature increased with both environmental temperature and quantity of food, indicating that the heat production associated with food intake adds to the heat load at high environmental temperatures. 5. Both heat production and abdominal temperature declined with decreasing light intensity and increased before feeding time. These effects were considered to result from changes in physical activity.     

The effect of rapeseed meal on the energy metabolism of laying hens

1. The effect of 100 g rapeseed meal (RSM)/kg diet on the energy metabolism of hybrid laying hens was examined by indirect calorimetry. Thyroid hormone concentrations, thyroid weight, liver weight and body weight, egg production and food intake were also measured. 2. Fasting heat production was significantly lower in hens receiving RSM than in controls, but this difference disappeared when the birds were fed. 3. Thyroid hormone concentrations decreased, while thyroid and liver weights increased slightly; none of these effects was significant. Body weight, egg production and food intake were unaffected and no liver haemorrhages were noted. 4. The maintenance metabolisable energy (ME) requirement of control and treated birds, estimated from short-term energy balance measurements, was 474 kJ/kg0.75 d; net availability of ME in both treatments was 0.85.     

Energy balance of laying hens selected on residual food consumption

1. Energy balance of adult hens divergently selected for high (R + ) or low (R- ) residual food consumption was investigated using indirect calorimetry. Three experiments were conducted: feeding behaviour of individual hens, hens having free access to food or fasting and hens tube-fed at 70, 100 or 130% of the control intake of both lines. 2. R + hens ate significantly more than R- (+ 48%). This difference was maximum at the onset of the light (+ 120%) and not significant during the rest of the light period. Although both lines spent the same total time eating, R + hens exhibited more frequent but shorter meals than R- ones, suggesting a higher feeding activity in R + hens. 3. True metabolisable energy (TME) intake was 28% greater in R+ than in R- birds. Basal heat production did not differ significantly between genotypes. Heat increment of feeding (HI) or diet-induced thermogenesis was significantly enhanced in R + birds: + 23.9 vs 13.7 kJ/100 kJ TME intake in R- . 4. When tube-fed and placed in darkness, restricted-fed R + hens had the same HI as R- birds. When fed at 100% and 130% of control intake, R + hens demonstrated a regulatory thermogenesis, which allowed them to dissipate the excess of energy ingested compared to R- hens. 5. No significant modification was observed in plasma triglyceride, phospholipid, uric acid, glucose or insulin concentrations between lines, suggestive of the use of similar thermogenic pathways. While plasma triiodothyronine (T3) concentrations did not differ between genotypes, plasma thyroxine (T4) appeared higher in unfed R + birds and equal in fed conditions in both lines. 6. A regulatory part of HI allowed R + hens to get rid of their excessive energy intake. An increased turnover of lipids could be involved.     

Effects of feather wear and temperature on prediction of food intake and residual food consumption

Heat production, which accounts for 0.6 of gross energy intake, is insufficiently represented in predictions of food intake. Especially when heat production is elevated (for example by lower temperature or poor feathering) the classical predictions based on body weight, body-weight change and egg mass are inadequate. Heat production was reliably estimated as [35.5-environmental temperature (degree C)] x [Defeathering (=%IBPW) + 21]. Including this term (PHP: predicted heat production) in equations predicting food intake significantly increased accuracy of prediction, especially under suboptimal conditions. Within the range of body weights tested (from 1.6 kg in brown layers to 2.8 kg in dwarf broiler breeders), body weight as an independent variable contributed little to the prediction of food intake; especially within strains its effect was better included in the intercept. Significantly reduced absolute values of residual food consumption were obtained over a wide range of conditions by using predictions of food intake based on body-weight change, egg mass, predicted heat production (PHP) and an intercept, instead of body weight, body-weight change, egg mass and an intercept.     

Peripheral ghrelin reduces food intake and respiratory quotient in chicken

Ghrelin injection, either centrally or peripherally strongly stimulates feeding in human and rodents. In contrast, centrally injected ghrelin inhibits food intake in neonatal chickens. No information is available about the mechanism and its relationship with energy homeostasis in chicken. Since ghrelin is predominantly produced in the stomach, we investigated the effect of peripherally injected ghrelin (1 nmol/100g body weight) on food intake and energy expenditure as measured in respiratory cells by indirect calorimetry for 24h in one-week-old chickens. Plasma glucose, triglycerides, free fatty acids, total protein and T(3) were measured in a separate experiment until 60 min after injection. Food intake decreased until at least 1h after intravenous ghrelin administration. The respiratory quotient (RQ) in ghrelin-injected chickens was reduced until 14 h after administration whereas plasma glucose and triglycerides concentrations were not altered. Free fatty acids and total protein levels also remained unchanged. Ghrelin did not influence heat production and this was supported by the absence of changes in plasma T(3) levels when compared to the control values. In conclusion, peripheral ghrelin reduces food intake as well as RQ and might influence the type of substrate (macronutrient) that is used as metabolic fuel.     

Preferred food rewards for laying hens in behavioural experiments

1. Laying hens showed a clear preference for maggots over whole grain wheat, feed pellets and other food stuffs in a preference test. 2. Motivation to work for maggots in an operant conditioning task was high and unaffected by periods of food deprivation of up to 6h before testing. 3. By extrapolation, motivation to work for maggots was equivalent to the motivation of the hens to work for their normal ration after approximately 9 h food deprivation. 4. Food deprivation is a common prerequisite for laying hens in operant conditioning and other behavioural techniques but can be avoided if an attractive food, such as maggots, is used. The mild distress associated with the deprivation is thereby removed and, in the UK, the procedure would not require regulation under the Animals (Scientific Procedures) Act 1986.     

The effect of egg formation and laying on the food and water intake of brown leghorn hens

Food and water intakes of four caged Brown Leghorn hens were studied on a daily and hourly basis in relation to egg formation. In addition the food intake of two similar hens was studied using a Skinner box which provided a more detailed record of ingestive behaviour in a non‐social situation in which social interactions might not mask the relationship between food intake and physiological needs. Food intake was greater on days on which ovulation occurred than on days during which there was neither ovulation nor oviposition. Water intake was greater on days during which ovulation occurred than on days with oviposition but no ovulation. On this latter type of day (laying day) food intake was greater than on days without ovulation and oviposition (resting day). Both food and water intakes were depressed for 1 to 2 h before oviposition, but ingestion increased during the hour of laying and remained high for 1 to 2 h.     

Food intake of the laying hen following crop loads of maize oil and other nutrients

The metabolisable energy (ME) of the diet of laying hens at an ambient temperature (Ta) of 20 degrees C was abruptly changed from 10.9 MJ/kg to 12.9 MJ/kg, or vice versa. Food intake during the next 14 d was significantly reduced by the low ME diet and was increased by the high ME diet, that is, the expected compensatory changes in food intake did not occur. Laying hens given the same change of diet as above but kept at 32 degrees C did not show any change in food intake within 14 d. Thus ME intake increased with the high ME diet and decreased with the low ME diet. Daily doses of 10 ml maize oil/kg body weight given directly into the crop of laying hens at a Ta of 20 degrees C, resulted in an immediate, significant, reduction of food intake such that total ME intake remained the same as with normal feeding. Daily doses of 3 ml maize oil/kg, given as before, resulted in an immediate, significant, reduction in food intake at a Ta of 20 degrees C but at a Ta of 32 degrees C food intake remained unchanged; consequently daily ME intake increased. Loading the crop with glucose or sucrose, at Ta 20 degrees C, in quantities which provided a similar ME as 3 ml maize oil/kg, reduced food intake but the adjustment was less precise and daily ME intake increased. Loading with glycerol or protein hydrolysate decreased both food and ME intake. Crop loads of starch were as effective as maize oil in bringing about a significant and compensatory reduction of food intake. Similar volumes of water or liquid paraffin placed in the crop did not affect food or ME intake. A similar weight or cellulose placed in the crop reduced food and ME intake.     

Effects of beak trimming and restraint on heart rate, food intake, body weight and egg production in hens

1. Heart rate (measured on restrained hens in two experiments) was used as an indicator of short term fear and pain responses of light and heavy strains of hens subjected to beak trimming. 2. In the first experiment 3 mm of the upper and lower mandibles was trimmed, while in the second 0, 2, 4, 6 and 8 mm of upper and lower mandibles were removed. 3. Production responses to beak trimming were measured after trimming, for 4 weeks in experiment 1 and for 10 weeks in experiment 2. 4. In the first experiment the recovery of beak trimmed hens to normal heart rate took significantly longer than that of control hens subjected only to catching and restraint, suggesting that there was short term pain associated with beak trimming. 5. The heavier strain took about 4 min longer to return to a normal heart rate than the lighter strain, indicating a strain difference in responsiveness to beak trimming. 6. Trimming the hens' beak by 3 mm had no significant effect on rate of lay or body weight, but their mean egg weight was depressed and food intake took 9 to 10 d to recover to pre-trimming values. 7. In the second experiment a plateau was reached in recovery time of the heart rate once 4 mm of beak was removed. Removal of 4, 6 and 8 mm of beak depressed normal feeding and resulted in variable effects on production and body weight.     

Influence of high dietary vitamin E supplementation on egg production and plasma characteristics in hens subjected to heat stress

1. The effects of different dietary concentrations of vitamin E (alpha-tocopherol acetate) were investigated in 2 experiments on laying hens exposed to chronic heat stress at 32C. 2. In the first experiment, egg production and plasma concentrations of calcium and egg yolk precursors were measured in 24 hens before, during and after a stress period of one week and fed on diets containing 10 or 500 mg vitamin E/kg. 3. In the second, larger experiment, egg production and food intake were measured in 300 hens housed in 2 temperature-controlled rooms and fed on diets containing 10, 125 or 500 mg vitamin E/kg. Birds in room 1 were stressed from 24 to 28 weeks of age and those in room 2 from 32 to 36 weeks. 4. In experiment 1, egg production and egg weight were significantly higher (72.6 vs 51.2%, P < 0.05 and 66.6 vs 63.1 g, P < 0.005 respectively) during and after the period of stress in the group given 500 mg vitamin E/kg. Plasma concentrations of calcium, vitellogenin (zinc) and VLDL (triglyceride) were also higher in this group. 5. In experiment 2, egg production was significantly higher (65.4 vs 56.2%, P < 0.05) during and after the period of heat stress in birds in room 1 fed on the diet containing 500 mg vitamin E/kg. Egg production was also higher (49.9% vs 44.7%) on this treatment during the stress period in room 2, though the difference was not significant (P < 0.10). Egg weight and food intake were unaffected by treatment in either room. 6. It is concluded that dietary supplementation with extra vitamin E can, at least in part, alleviate the adverse effects of chronic heat stress in laying hens, perhaps by maintaining the supply of egg precursors in plasma.     

Maintenance energy requirements of laying hens: a comparison of measurements made by two methods based on indirect calorimetry

1. The aims of this study were to compare maintenance metabolisable energy requirement estimates obtained in laying hens by two methods of respiration calorimetry, to investigate the changes in the heat production and energy retention of laying hens associated with temporarily reduced energy intake and to demonstrate the errors caused by using inappropriate values of net efficiency of energy utilisation (k) when extrapolating to maintenance energy requirement from a range of energy retention values. 2. One of the methods used regression of a range of energy retentions on energy intakes, followed by interpolation of zero energy retention; the other method employed extrapolation to zero energy retention from a single level of energy intake and retention. 3. Maintenance energy requirements derived from the two methods were similar. There was no indication that repeated temporary reduction of energy intake for the regression method gave a reduced value for maintenance energy requirement. The main potential advantage of the single-point method is therefore rapidity. 4. The single-point method relies for accuracy either on fixing an energy intake near the expected maintenance or on having a very accurate value for k. For example, the error attributable to using an assumed net efficiency (k) value of 0.7 when the true value was 0.8 was shown to vary between 0 if energy retention was measured at the equilibrium point and about 70 kJ/kgW0.75 d, or +15%, if energy retention was measured in the fasted state. 5. Stability of heat production and energy intake was approached after 2 d of reduced energy intake.     

Effect of excess dietary sodium, potassium, calcium and phosphorus on excreta moisture of laying hens

1. Four experiments were conducted to investigate the effects of dietary concentrations of sodium, potassium, calcium or phosphate on the water intake and excreta moisture of laying hens. A fifth experiment examined the effect on these variables of increasing amounts of 2 different sodium salts (chloride or bicarbonate) and the interactions with 2 levels of dietary phosphorus. 2. All experiments involved individually caged laying hens fed on diets varying in 1 or 2 minerals in replacement for washed sand. The experimental diets contained mineral concentrations that either met or exceeded the expected requirement of the hens. The diets were given for a 7 or 8 d feeding period and food and water intakes were measured and excreta were collected for the last 48 h of each feeding period. These data were corrected for evaporative water loss to the environment during the collection period. 3. Increasing dietary concentrations of sodium, potassium or phosphorus gave linear increases (P<0.001) in the water intake of the laying hens and linear increases (P<0.01) in the moisture content of their excreta. Each 1 g/kg increase in dietary mineral increased the moisture content of the excreta by 9.04 (+/- 1.57), 11.95 (+/- 2.02) and 5.59 (+/- 0.31) g/kg (+/- standard error) for sodium, potassium and phosphorus, respectively. Increasing concentrations of dietary calcium did not significantly affect the water intakes or excreta moisture levels of the laying hens. 4. The fifth experiment showed that, although there was a sodium x phosphorus interaction (P<0.05), the effects of the 2 mineral additions were approximately additive. There were no significant differences (P>0.05) in water intakes or excreta moisture contents due to the 2 different sodium salts (chloride or bicarbonate).     

Effects of environmental temperature and heat production due to food intake on abdominal temperature,shank skin temperature and respiration rate of broilers

1. Eight broilers were used to determine the effects of environmental temperature and the increased heat production attributable to food intake on thermoregulatory physiological responses. Heat production, abdominal temperature, shank skin temperature and respiration rate were measured.

2. Heat production rose with increase in food intake and environmental temperature. Abdominal temperature, shank skin temperature and respiration rate also increased but, at 36°C, there was no difference in respiration rate between the different rates of food intake after exposure for 1 h. Also, heat production decreased with exposure time when the birds were exposed to 28°C and 32°C, but increased at 36°C ambient temperature.

3. Although the effect of the increased heat production on thermoregulatory physiological responses was not greater than that of environmental temperature, the increase in heat production enhanced the effect of ambient temperature on thermoregulatory physiological responses. However, with increasing heat production, the greatest response in shank skin temperature was at 28°C, in respiration rate at 32°C and in abdominal temperature at 36°C.

4. These results suggest that, at high environmental temperatures, arranging feeding management to restrict the increase in heat production may alleviate the effect of environmental temperature.     

Effects of lighting regimen and grower diet energy concentration on energy expenditure,fat deposition and body weight gain of laying hens

1. Mean metabolisable energy (ME) intakes and heat productions over a laying year were calculated for laying hens which had been submitted to one of various lighting regimens and given either a normal or a high energy ration during the rearing period.

2. Daily ME intake and heat production per hen in the laying period were unaffected by either lighting regimen or grower diet. ME intake per kg W ^0.75 and heat production per kg W ^0.75 during lay increased significantly with laying photoperiod, was non‐significantly higher following an 8‐h rather than an 11‐h rearing photoperiod, but was unaffected by dietary energy concentration. The increase in heat production (/kgW°‘⁷⁵) associated with a 1‐h increment in photoperiod was similar to predictions made from calorimetric measurements of diurnal variation.

3. Efficiency of conversion of food to egg was unaffected by either lighting regimen or dietary energy concentration.

4. Fat weight gain in lay was not influenced by lighting regimen, but was significantly lower in birds reared on the high, compared to the normal, energy grower ration. Fat‐free weight gain in lay was unaffected by grower diet, but was significantly increased by photoperiods longer than 8 h.

5. ME intake and heat production per kgW^⁰⁷⁵ were negatively correlated with age at first egg, but ME intake and heat production per bird d were not related to age at sexual maturity.     

Influence of dietary energy, nutrient density and environmental temperature on pullet performance in early lay

1. Alterations in dietary metabolisable energy (ME) concentration had a limited influence on food and nutrient intakes and egg mass output of hens in early lay kept at the prevailing air (10 degrees to 24 degrees C), cold (6 degrees to 16 degrees C) or hot (25 degrees to 35 degrees C) temperatures. 2. Energy intakes were not improved by increasing the dietary concentrations of nutrients other than energy. 3. At prevailing air and cold temperatures all dietary ME-nutrient density combinations allowed hens to meet the recommended daily protein intake but only hens fed the most concentrated diets were able to meet this recommendation at hot temperatures. 4. Even the highest intakes of ME and protein achieved at hot temperatures failed to increase egg mass output to the values attained on any diet at cold temperatures.     

Effects on egg production of including high residual lipid copra meal in laying hen diets

1. The effects on egg production of including copra meals of 220.9 (Copra‐1) and 205.0 (Copra‐2) g residual lipids/kg, in laying hen diets at up to 400 g/kg, were examined in two feeding trials. In a separate experiment, the dry matter retention and ME of a 400 g Copra‐2/kg diet were determined between 6 to 9, 9 to 12 and 12 to 15 d of feeding.

2. Food intakes, egg output and efficiencies of food utilisation (g eggs/g food) of hens fed on Copra‐1 at 100 and 200 g/kg diet were not significantly different from controls; while food intakes at 400 g Copra‐1/kg diet were similar to controls, egg output (6.0% lower), efficiency of food utilisation and body weights of hens were significantly reduced. Hens fed Copra‐2 at 400 g/kg diet had 5.0 and 6.9% lower food intakes and egg output, respectively, and lost body weight compared to controls; however, egg output increased to that of the control group by the 11th week of feeding.

3. Initially, dry matter retention of the 400 g Copra‐2/kg diet was similar to, and ME slightly higher than the control diet; however, both were reduced significantly in the second and third collection periods. The water content of excreta from hens fed on 400 g Copra‐2/kg diet was significantly higher than that from controls.

4. Complete replacement of dietary maize, mainly with copra (experiment 1) led to an increase in pale yolks, with the 400 g Copra‐1/kg diet producing cream‐coloured yolks. Inclusion of maize at 180.6 g/kg of the 400 g Copra‐2/kg diet (experiment 2) resulted in eggs with slightly golden‐yellow yolks.     

Influence of a high-protein diet on energy balance in obese cats allowed ad libitum access to food

The influence of a high-protein [HP, 47% of metabolizable energy (ME)] diet on energy balance was evaluated in obese cats allowed ad libitum access to food. Energy intake, body weight, body composition, energy expenditure, and concentrations of hormones and metabolites associated with carbohydrate and lipid metabolism (glucose, insulin, free fatty acids, triglycerides and leptin) were measured in cats after consuming either a moderate protein (MP, 27% of ME) or HP diet for 4 months. Indirect respiration calorimetry showed that resting and total energy expenditure (kJ/day) adjusted for either body weight or lean body mass was increased in cats consuming the HP in relation to MP diets. However, voluntary energy intake also was increased in the HP treatment and, thus, there was no difference in body weight between animals consuming the two diets. Body composition measurements using deuterium oxide dilution showed that dietary protein content did not alter amounts of either lean body mass or fat mass. No significant differences (p > 0.05) were observed between the two treatment groups for blood glucose, free fatty acid or leptin concentrations, although there was a trend (p = 0.054) towards an increase of serum insulin concentrations in the cats eating the HP diet. This study showed that short-term ad libitum feeding of an HP diet did not reduce food intake or promote weight loss in obese cats. However, energy expenditure was increased in the HP diet group and it is possible that this effect of HP might help promote weight loss when energy intake is restricted.     

Effect of diurnal or nocturnal heat stress on egg formation.

1. Mature laying hens were subjected to ambient temperatures sufficient to maintain body temperature of 43 degrees C for periods of 6 to 7 h during the day (eight periods) or the night (five periods). This did not reduce total daily food consumption. 2. The effect of heat stress during the day was mostly on egg-shell quality, being adverse and significant. During the night heat stress caused a significant decrease in egg production, its effect during the day was less marked in this respect. 3. Heat stress thus seems mostly to affect the early stages of shell formation, while its effect on egg production seems to depend on the time at which it occurs in relation to ovulation. 4. These results suggest that egg production is directly affected by heat stress; prolonged heat stress probably also acts indirectly, by suppression of food intake.     

Effects of food intake regulation on the energy metabolism of hens and cockerels of a layer strain

1. Indirect calorimetry was used to study the effects on the energy metabolism of cockerels and hens after peak lay of a hybrid layer strain of regulating food or energy intake to 80% of ad libitum for an extended period of time. 2. Regulation of food intake decreased fasting and fed heat productions per bird and per unit metabolic body weight (kg0.75). 3. Maintenance energy requirements (per kg0.75) of both sexes were reduced about 20% by regulation; maintenance requirements of cockerels were about 30% lower per kg0.75 than hens on the same treatments. 4. Reducing the intake of all nutrients by 20% had the same effect on the energy metabolism of cockerels as restricting energy alone by the same amount. 5. Gross efficiency of egg production, in terms of both mass and energy, was higher when intake was regulated. During weeks 21 to 52 of restriction, daily egg mass output was significantly higher in hens fed ad libitum. 6. After 52 weeks of food regulation, restricted hens weighed 20% less and their total carcass energy was 36% less than hens fed ad libitum; similar effects were found in the cockerels.