Why cuckoos should parasitize parrotbills by laying eggs randomly rather than laying eggs matching the egg appearance of parrotbill hosts?

The coevolutionary interaction between cuckoos and their hosts has been studied for a long time, but to date some puzzles still remain unsolved. Whether cuckoos parasitize their hosts by laying eggs randomly or matching the egg morphs of their hosts is one of the mysteries of the cuckoo problem. Scientists tend to believe that cuckoos lay eggs matching the appearance of host eggs due to selection caused by the ability of the hosts to recognize their own eggs.In this paper, we first review previous empirical studies to test this mystery and found no studies have provided direct evidence of cuckoos choosing to parasitize host nests where egg color and pattern match. We then present examples of unmatched cuckoo eggs in host nests and key life history traits of cuckoos, e.g. secretive behavior and rapid egg-laying and link them to cuckoo egg laying behavior. Finally we develop a conceptual model to demonstrate the egg laying behaviour of cuckoos and propose an empirical test that can provide direct evidence of the egg-laying properties of female cuckoos. We speculate that the degree of egg matching between cuckoo eggs and those of the host as detected by humans is caused by the ability of the hosts to recognize their own eggs, rather than the selection of matching host eggs by cuckoos. The case of Common Cuckoos(Cuculus canorus) and their parrotbill hosts(Paradoxornis alphonsianus), where it has been shown that both have evolved polymorphic eggs(mainly blue and white), was used to develop a conceptual model to demonstrate why cuckoos should utilize parrotbill hosts by laying eggs randomly rather than laying eggs matching the appearance of host eggs.In conclusion, we found no evidence for the hypothesis that cuckoos lay eggs based on own egg color matching that of the parrotbill-cuckoo system. We argue theoretically that laying eggs matching those of the hosts in this system violates a key trait of the life history of cuckoos and therefore should be maladaptive.     

Phosphorus requirements of laying hens fed on wheat‐based diets

1. Three experiments were carried out to determine the phosphorus (P) requirements of laying hens aged 34 to 58 weeks (experiment 1), 59 to 70 weeks (experiment 2) and 22 to 50 weeks (experiment 3) given diets containing wheat, sorghum and soyabean meals as the main ingredients. Dietary total P (P_t) varied between 3.2 and 7.3 g/kg (experiment 1), 3.2 and 4.6 g/kg (experiment 2) and 3.0 and 6.6 g/kg (experiment 3). Hens were housed at either 18° or 30°C (experiments 1 and 2) and uncontrolled temperature (experiment 3), and in experiment 2 diets were fed without or with a phytase supplement of 500 units/g.

2. Dietary P_t had no significant effect on production measures in any experiment. Increases in dietary P_t adversely influenced egg shell quality although uterine calcium (Ca), ATPase and carbonic anhydrase activities were unaffected.

3. A 3‐d‐feeding trial in experiment 1 gave maximum P_t retentions of 228 mg/d at 18°C and 204 mg/d at 30°C. These were obtained with diets containing, respectively, 4.6 and 6.0 g P_t/kg.

4. Plasma inorganic P (P_i) increased consistently with increases in dietary P_t at all temperatures but plasma total Ca, and tibia Ca and P, were unaffected.

5. The inclusion of the phytase supplement in diets containing 3.2 and 4.6 g P_t/kg had an adverse effect on egg production at both temperatures in experiment 2.

6. A dietary P_t concentration of 3.2 g/kg, providing a calculated 1.2 g available P (P_av) /kg, with a dietary phytase activity of less than 200 units/kg, satisfied the P requirements of the hens used in these studies. However, the data from experiment 3 suggest that the P_t requirement of some flocks fed on wheat‐based diets may be lower than 3.2 g/kg.     

Effects of lauryl sulphate on egg‐shell quality in laying hens

1. The effect of feeding the detergent, sodium lauryl sulphate at concentrations up to 10 g/kg diet on egg‐shell quality was examined in laying hens at 24, 36 and 48 weeks of age.

2. Lauryl sulphate at 2.5 g/kg or more in the diet significantly increased shell breaking strength, shell thickness and shell weight at 24 and 36 weeks. In 48‐week‐old hens only shell breaking strength was increased significantly. Egg production and egg weight were not affected significantly by lauryl sulphate.

3. Lauryl sulphate might increase these measures of quality by increasing shell deposition and/or shell elasticity.     

A test for photorefractoriness in high‐producing stocks of laying pullets

1. Pullets of 2 high‐producing commercial stocks (both brown‐egg layers) were exposed to 5 different lighting patterns between 18 and 72 weeks to test the hypothesis that photoperiods used in commercial lighting programmes early in the laying year may be unnecessarily long and, by accelerating the development of photorefractoriness, may contribute to the decline in egg production observed after the initial peak. Two rooms of 288 pullets were allocated to each treatment.

2. The rate of lay observed with a Step‐Up treatment which gave increases in photoperiod from 8L:16D at 18 weeks to 15L:9D at 27 weeks of age was not significantly different from that of treatments which held the birds on 11L:13D during peak egg production but gave increments up to 15L:9D later in the laying year.

3. A control group maintained on 11L:13D from 20 to 72 weeks laid 295 eggs per bird housed and a further group held on 8L:16D from 0 to 72 weeks laid 284 eggs per bird. These yields were lower than the Step‐Up treatment (299 eggs) but show the potential of modern hybrid stocks to lay prolifically even without light stimulation.

4. It is concluded that the stocks tested in this experiment showed no advantage when given lighting programmes in the first laying year which were designed to minimise the adverse effects of photorefractoriness.     

Responses of the reproductive vascular system during the egg‐formation cycle of unanaesthetised laying hens

1. Vascular adjustments to the process of egg formation were examined in the unanaesthetised laying hen, by the radioactive microspheres method.

2. Three‐ to four‐fold increases in blood flow were found in segments surrounding the egg during its passage along the oviduct, possibly due to an enhanced metabolic activity in the muscle layer of the oviduct.

3. Shell‐gland blood flow was minimal in the absence of an egg and increased gradually to a maximum (5‐fold) about 5 h after entrance of the egg into the shell gland. This parallels the rate of calcification of the egg shell.

4. Changes in blood flow in the ovarian follicles and other parts of the oviduct were small while the egg was in the shell gland. This might be typical for the reproductive system of the fowl, which undergoes little structural alteration during egg formation.     

The egg‐laying response of the domestic hen to variation in dietary calcium

1. Eight diets (1 to 8) containing respectively 36.8, 30.9, 19.6, 9.7, 5.5, 1.9, 0.96 and 0.48 g of calcium/kg food were each fed for 11 weeks to 24 individually‐caged laying hens (Gallus domesticus) aged 24 weeks. After the experimental period, all birds were returned to a normal diet (34.9 g calcium/kg).

2. Food consumption and egg production decreased as dietary calcium decreased. Shell weight was unaffected on diets 1 and 2; on diet 3 there was slight reduction of shell weight and on diets 4 to 8 the reduction was marked. The proportion of calcium in the shell was affected particularly on diets 7 and 8, though those from diet 5 also showed a decreased shell calcium.

3. The values for calcium intake and calcium loss in the egg showed that, generally, birds restricted calcium loss to less than intake. Only on the very low concentrations of calcium (diets 6, 7 and 8) did output appear to exceed input.

4. The main mechanism for controlling calcium loss involves the regulation of the number of eggs produced, i.e. the number of ovulations. Alterations in shell quality are of less importance with respect to calcium balance, although shell strength was impaired on the more restrictive diets (5 to 8).     

Disease outbreak in laying hens arising from the consumption of fumonisin‐contaminated food

1. During the last quarter of 1995 an unusual disease outbreak characterised by black sticky diarrhoea, severe reduction in food intake, egg production and body weight follow by lameness and death was observed in 2 layer farms in Andhra Pradesh. A total of 6700 hens of 64 weeks age and 3000 hens of 36 weeks age were affected.

2. Around 10% mortality and a 20% reduction in egg production was observed. Postmortem examinations showed pale yellow coloured livers with peripheral congestion, mild haemorrhage in the proventriculus and watery accumulations in the intestine.

3. The disease was traced to a new batch of food and its withdrawal improved the health status of the flocks.

4. Analysis of the diet indicated contamination with fumonisin B₁ up to 8–5 mg/kg and with aflatoxin B₁ up to 0.1 mg/kg.

5. Diarrhoea was induced in day old cockerels by feeding the suspect diet containing 8.5 mg/kg fumonisin Bi, and in laying hens by feeding a normal diet with fumonisin B₁ additions of 8 and 16 mg/kg.     

Ultrastructural changes of osteoclasts on hen medullary bone during the egg‐laying cycle

1. The ultrastructure of osteoclasts on hen medullary bone during the egg‐laying cycle was observed by electron microscopy.

2. At 0 to 6 h after oviposition, osteoclasts lacking ruffled borders were attached to the bone via the clear zone and appeared to have ceased bone resorption. Small vacuoles were scattered throughout this cytoplasm.

3. At 9 to 21 h after oviposition, most of the osteoclasts had ruffled borders and appeared to be resorbing bone. The ruffled borders at 15 h were well developed, whereas at 9, 18 and 21 h they were poorly developed or showed similarities in structure to the clear zone. The small vacuoles were concentrated under the ruffled borders at 9 h and then decreased.

4. These results demonstrate cyclic changes in osteoclasts during the egg‐laying cycle, indicate that ruffled borders form at the beginning of bone resorption and suggest that they are derived from the clear zone and the small vacuoles. The results also indicate that the ruffled borders fuse into the clear zone and disappear at the completion of bone resorption.     

Growth and laying performance of light‐hybrid pullets subjected to quantitative food restriction

1. Three degrees of quantitative food restriction producing body weights 88, 76 or 64% of controls at 20 weeks and three times of starting restriction 3, 6 or 9 weeks of age were compared with ad libitum‐fed pullets in a factorial experiment with 1800 light‐hybrid hens. Restrictions were applied such that the target weights were produced irrespective of time of starting.

2. Food intake was least when restrictions were started at 3 weeks.

3. Sexual maturity was delayed in proportion to the severity of restriction and resulted in lower percentages of small eggs.

4. Egg numbers decreased as restriction increased.

5. Mortality in restricted groups during rearing and laying was higher.

6. Optimal restriction was apparently intermediate between treatments producing 88 or 76% of ad libitum‐fed body weight. Body weight was so excessively affected by the most severe restriction that total production was adversely affected.

7. Restriction from 3 weeks was more profitable than restriction from 6 or 9 weeks, resulting in lower food cost during rearing and higher total egg production.     

Effect of selected dietary antioxidants on fatty liver‐haemorrhagic syndrome in laying hens

1. Single comb White Leghorn hens of an inbred line highly susceptible to fatty liver haemorrhagic syndrome (FLHS) were fed supplemented dietary ascorbic acid (200 mg/kg), α‐tocopherol (75 mg/kg), or L‐cysteine (3 g/kg, and 6 g/kg) for 28 d in order to evaluate the potential therapeutic effect of these compounds against the disease.

2. Supplementation of ascorbic acid, α‐tocopherol, or a low level of L‐cysteine (3 g/kg) did not significantly affect any of the hepatic variables evaluated. Hepatic glutathione was not increased by the supplementation of dietary L‐cysteine.

3. L‐cysteine supplemented at a level of 6 g/kg decreased hepatic dry matter and fat contents without affecting the hepatic malondialdehyde or the liver haemorrhagic score.

4. Because one of the predisposing factors of FLHS is a high hepatic fat content it was concluded that dietary supplementation of L‐cysteine (6 g/kg) may be useful in the prevention of the disease.     

Battery gage shape: The laying performance of medium‐ and light‐body weight strains of hens

1. One thousand two hundred and ninety‐six medium‐ and 1296 lightweight hens were housed, four to a cage, in deep (conventional) cages, 405 mm wide by 460 mm deep, or shallow cages, 610 mm wide by 305 mm deep, between 18 and 70 weeks of age. The performance of the hens during different periods of the laying stage and over the full laying year was measured.

2. Egg number per medium‐weight hen housed in shallow cages was significantly higher than of hens housed in deep cages between 18 and 30 weeks and 31 and 42 weeks. Thereafter significant differences were not found. With the light‐weight hens the increase was significant only in the 18‐ to 30‐week period. Apart from the 19‐ to 30‐week period hens of both strains housed in shallow cages consumed significantly less food than those in deep cages, the reduction over the full year being about 4%. In all periods the hens housed in the shallow cages converted food more efficiently. Gage shape had no significant effect on mortality.

3. The incidence of hair‐cracked, cracked and broken eggs between 60 and 70 weeks of age from hens housed in shallow cages was lower than that from hens housed in the deep cages but only with the light‐weight hens did this difference reach significance. Shell thickness, shell weight and albumen quality of eggs laid by hens housed in the different shaped cages were not significantly different.     

Individual and combined effects of t‐2 toxin and das in laying hens

1. The individual and combined effects of T‐2 toxin and 4,15‐diacetoxyscirpenol (DAS) on laying hens were investigated in an experiment consisting of a 2 X 2 completely randomised factorial design with dietary concentrations of 0 and 2 mg/kg T‐2 toxin and 0 and 2 mg/kg DAS.

2. Individually, T‐2 toxin and DAS induced oral lesions in half of the hens and decreased significantly egg production and food intake.

3. The effects of T‐2 toxin and DAS were additive for reduced food consumption and incidence of oral lesions. However, a synergism for reduced egg production was observed during the last experimental period.

4. No effects on body weight were observed during this study. Mild changes in selected plasma enzymes activities and no change in liver malondialde‐hyde content were detected.

5. The combination of T‐2 toxin and DAS was more toxic than the single mycotoxins, for some parameters, and therefore, may pose a greater economic threat to the poultry industry than either of the toxins individually.     

High,low, or familiar? Nest site preferences of experienced laying hens

1. The aim of this study was to investigate which nest heights are preferred by laying hens in the absence of familiar nest locations and whether preferred nest heights are more attractive than a familiar location. In two experiments, a total of 108 hens of four different layer breeds, which were at least 50 weeks of age, were studied.

2. In the first experiment, hens were given individual free choice between nests for 1-week at four different heights (0 cm, 39 cm, 78 cm, and 117 cm above ground). Hens of the four breeds differed in their nest height preferences (P = 0.0013). However, hens of three breeds preferred ground level nests (P < 0.007) and the fourth line showed an equal preference for the ground level and level three, the latter level corresponding to the height of the nests in their home compartments.

4. In the second experiment, hens from the four breeds were given a choice between ground level nests and nests at a familiar location, i.e. at the same location as in their home compartment. Hens of all strains preferred the familiar nest location (P = 0.002) and preferences did not differ between strains (P = 0.77).

5. Laying hens seem to prefer nests at ground level in the absence of a familiar nest. However, if possible, experienced 50 week old hens continue to use a familiar nest location instead of a ground nest location. The results are discussed with respect to a potential primary preference that may be modifiable by experience and with respect to possible relevance in commercial housing.     

The effect of sodium sulphate and methionine on the utilisation of non‐protein nitrogen by laying hens

1. The response of layers to adding diammonium citrate (DAC), sodium sulphate or methionine to a basal diet containing 136–3 g protein/kg was determined.

2. Supplementing the diet with DAC equivalent to 25 g protein/kg did not improve egg production, the efficiency of food utilisation, egg weight, nitrogen retention or the apparent absorption of lysine and methionine; increases in food intake and in the concentration of methionine in the serum and liver were observed.

3. Adding sodium sulphate, alone or with DAC, did not affect the variables noted above.

4. Supplementation of the basal diet with methionine increased egg production, egg weight, food intake and the concentrations of lysine in the serum and liver.

5. It is concluded that the supplemental NPN was used only in serum protein synthesis.     

Comparative analysis of pre‐laying behaviour in battery cages and in three alternative systems

1. Pre‐laying behaviour was compared in battery cages (C) and in three alternative husbandry systems: perchery (P), deep‐litter (DL) and covered strawyard (S).

2. Twenty ISA Brown hens were observed in each system for 60 min before oviposition and 5 min afterwards.

3. There were significant between‐system differences in a number of behavioural elements associated with the nest. Most varied along the sequence: C, P, DL, S, but the differences between the three alternative systems were generally less than between them and C.

4. The main factor governing the expression of pre‐laying behaviour appeared to be the availability, the number and the arrangement of littered nests.

5. Behaviour elements preceding nest entry and which were associated with area per bird again varied along the sequence C, P, DL and S with the major discontinuity between P and DL.

6. In P and C the searching phase was extended, in C at the expense of sitting. In P much time was spent inspecting the nests and many nest entries were made.

7. The Gakel call appeared to be related to finding a suitable nest site. In P the number of calls and the mean duration per nest inspection declined in a parallel fashion towards oviposition. In C, where no suitable nest could be found, Gakel calling remained high during the whole observation period.

8. These behavioural differences suggested that pre‐laying behaviour was poorly expressed in C and most fully in S, but that expression was in general relatively complete in all three alternative systems.

9. The ability of hens to express pre‐laying behaviour fully and coherently may be relevant to their welfare.     

A formal method of determining the dietary amino acid requirements of laying‐type pullets during their growing period

1. The amino acid requirements of laying type pullets during the growing period can be estimated by measuring the growth of different components of the body and making use of nutritional constants that define the amount of each amino acid that is required for the production of the tissues being formed.

2. In this experiment, carcase analyses of each of three breeds of pullets were conducted at weekly intervals throughout the growth of the pullets, to 18 weeks of age. Measurements were made of body weight, gut‐fill and feather weight, and chemical analyses consisted of water, protein, lipid and ash measurements of both the body and the feathers. Each age group comprised 10 birds of each breed.

3. Gompertz functions accurately estimated the growth of both body protein and feather protein, to 18 weeks of age, from which the rate of growth of these two components of the body could be estimated. The mature weight of pullets was overestimated by the Gompertz growth curve, which may indicate that a pullet ceases to increase in body protein content once sexual maturity has been reached.

4. Using allometric relationships between the chemical components of the body and of feathers, all the components of growth could be estimated from the growth of body protein and feather protein. These components were then added together to determine the growth rate of the body as a whole.

5. The daily amino acid requirements for 4 functions were calculated, namely, those for the maintenance of body protein and feather protein, and for the gain in body protein and feather protein. These requirements were then summed to determine the requirement of pullets on each day of the growing period.

6. Using the ‘effective energy’ system, the amount of energy required by these pullets was calculated for each day of the growing period, from which the desired daily food intake of the pullets could be predicted. By dividing the amino acid requirement by this daily food intake it was possible to determine the concentration of amino acids that would be needed in the diet in order to meet the requirements of a pullet.

7. The results indicate that the ratio between the requirement for lysine and for methionine and cysteine changes dramatically during the growing period, negating the concept of a fixed ratio between all the amino acids during growth.

8. The above process is the first step in determining the optimal feeding programme for a population of pullets of a given genotype. The constraining effects, of the diet being offered and of the environment in which the pullets are housed, on the food intake and growth rate of each pullet have to be estimated, and such a theory can then be expanded to include all the individuals in the population. Only by the use: of simulation models can all these constraining effects be considered simultaneously.     

Feeding value of rice distiller’s dried grains with solubles as protein supplement in diet of laying hens

A feeding trial of 10 weeks duration was undertaken on laying hens (n = 240) to evaluate feeding value of rice distiller’s dried grains with soluble (rDDGS) with or without enzyme supplementation (α-amylase, β-glucanase, xylanase, carboxymethylcellulase, pectinase, proteinase, α-galactosidase, β-galactosidase, lipase, and phytase), following 4 × 2 factorial design, on egg production, nutrient utilization, and cost economics of egg production. The birds were randomly assigned to eight dietary treatments with 30 birds/treatment. The birds were housed individually in layer cages and each bird was taken as an experimental unit. Eight experimental diets were prepared by incorporating four levels (0, 50, 75, and 100 g/kg) of rDDGS with and without enzyme supplementation. The results revealed a significant (P < 0.01) increase of egg mass, feed intake, egg production, and body weight gain in dietary treatments with up to 75 g rDDGS though the values were statistically similar to the hens fed 100 g rDDGS. Enzyme supplementation resulted in significant (P < 0.01) improvement of egg mass, egg production, feed conversion ratio (FCR) per dozen eggs, FCR per kilogramme egg mass, and net FCR. The significantly (P < 0.01) higher yolk index was observed at 100 g rDDGS level, while shell thickness improved significantly (P < 0.01) up to 75 g rDDGS level. No significant effect of rDDGS inclusion was observed on shape index, albumin index, and Haugh unit. Enzyme supplementation significantly improved the shell thickness and yolk colour of eggs. Nitrogen, calcium, and phosphorus retention and dry matter metabolizability did not show any significant treatment effects. There was significant (P < 0.01) reduction in feed-cost per kilogramme egg mass or per dozen eggs with the increased DDGS levels and dietary enzyme supplementation. It was concluded that rDDGS can be used up to 100 g/kg diet of laying hens along with enzyme supplementation for better productivity of layer hens.

     

Effects of plastic anti‐pecking devices on food intake and behaviour of laying hens fed on pellets or mash

1. This study investigated effects of plastic anti‐pecking devices on food intake and behaviour to assess their applicability in commercial layer flocks.

2. In experiment 1, 72 individually caged ISA Brown pullets were fitted at 16 weeks of age (point of lay) with either ‘ring’ or ‘bumper’ beak devices or no device (control), half being fed on pellets and half on mash. The devices were held in place by lugs inserted in the nares.

3. There was evidence of discomfort immediately after fitting the devices, but not thereafter. In the first week, the devices reduced food intake, and were removed from 7 (15%) birds that lost weight consistently. Between 16 and 21 weeks, however, there were no significant effects of beak treatment or food form on either total food intake or egg production with the 65 remaining birds.

4. Beak treatment did not affect feeding efficiency (food intake per minute of feeding). The only behaviour affected was pecking at birds in adjacent cages, which was reduced with the ring device.

5. At 21 weeks, the birds were placed together in groups of 4 to 6 in 12 pens, each group consisting of birds from one of the 6 original treatments, and their behaviour was observed in experiment 2 between 21 and 23 weeks.

6. The only effects of beak treatment on behaviour were that pecking at pen walls was increased with the ring device, while pulling and eating feathers from group mates was seen mainly with control birds fed on pellets. Two birds were cannibalised in a control group on pellets.

7. Devices came off 3 (7%) birds in experiment 1 and were replaced, and rings slipped over lower mandibles of 2 (5%) birds in experiment 2 and were corrected.

8. At 23 weeks, beak lengths were increased with ring and bumper treatments, and pecking damage scores were low with all treatments.

9. It is concluded that these devices are not applicable commercially, but further trials would be justified with new improved designs.