Local cutaneous water barrier in cold- and heat-acclimated pigeons (Columba livia) in relation to cutaneous water evaporation

The thermoregulatory function of the skin differs in adult cold-acclimated and heat-acclimated rock pigeons (Columba livia). In general, the cutaneous evaporative cooling mechanism is not activated by appropriate stimuli in cold-acclimated pigeons in contrast to heat-acclimated pigeons. We studied with electron microscopy whether the differences in the function of the skin are reflected in the structure of the epidermal water barrier of these two extreme acclimation states. The epidermis of cold-acclimated pigeons is attenuated, and the underlying dermis lacks any intimate vascularization. Both the extracellular and the intracellular domains in the stratum corneum contain organized lamellar lipids. At the stratum transitivum-stratum corneum interface, multigranular body secretion is indicated by the highly convoluted cell membranes and membraneous sacculae enclosing the multigranular bodies. Alternatively, multigranular bodies retain in the corneocytes, and the lipoid material originated from them is reprocessed to broad lamellae. The keratohyalin (KH) granules are spotlike and oriented as cortical bands beneath the plasma membrane. In heat-acclimated pigeons, the epidermis displays modified patches side by side with basic structural type of epidermis. The modified areas are characterized by hypertrophy and abundance of dermal capillaries adjacent to the hypertrophied patch. No lamellar lipids are discerned in the dilated extracellular space. The structure of multigranular bodies is abnormal, and the numbers of lipid droplets in the outer viable epidermis and stratum corneum are decreased. The transitional cells contain stellate KH granules, which form a network throughout the cell. It is concluded that cold-acclimated pigeons have a lamellar, extracellular water barrier, the cutaneous water evaporation is minimized, and heat is stored in the body core. Acclimation to heat leads to formation of structurally heterogeneous skin. The structurally modified skin patches show disruption of the barrier-forming machinery in the multigranular bodies and marked reorganization of fibrillar proteins and electron-dense KH masses in the transitional layer. Thus water barrier adjustments in cold- and heat-acclimated pigeons manifest the dynamic function of avian skin as a thermoregulatory organ.     

Cooling by cutaneous water evaporation in the heat-acclimated rock pigeon (Columba livia)

The present study provides an up-to-date overview of the cutaneous water-evaporation cooling mechanism in the rock pigeon. Cutaneous water evaporation fully replaces the classic respiratory cooling mechanism in the resting, heat-acclimated bird, and is more economical in terms of water conservation. It enables the pigeon to maintain homeostasis, and to breed successfully in harsh environments. Adrenergic signaling is involved in the initiation of this novel mechanism, either by deactivation of the beta-adrenergic receptors (ARs), or activation of the alpha-AR. The adrenergic signaling results in a marked increase in cutaneous blood flow and in the arterial-to-venous blood-flow ratio. This is associated with alterations in the cutaneous capillary wall ultrastructure, which increase its permeability to plasma proteins and water. The end result of this process might be an increase in water efflux from the capillary lumen. The properties of beta-ARs were measured in the cardiac muscle of thermal-acclimated pigeons. Significant down-regulation in the density of beta-ARs, associated with increased affinity of these receptors, was measured in the heat-acclimated pigeon. Concomitantly, changes in the skin ultrastructure and lipid composition were found in very well defined patches in the epidermis of heat-acclimated pigeons. These suppress the skin resistance to water transfer. We suggest that this cooling mechanism involves finely orchestrated adjustments in the ultrastructure of the skin and the cutaneous capillaries, and in skin blood flow. Adrenergic signals are among those factors that regulate this cooling mechanism during exposure to a hot environment.     

Hepatic and extra-hepatic glutathione-S-transferase activity in wild pigeons (Columba livia)

Glutathione-S-transferase (EC 2.5.1.18) activity was assayed in hepatic and extra-hepatic tissues of pigeons using l-chloro-2,4-dinitrobenzene and 1,2-dichloro-4-nitrobenzene as substrates. Gluthathione-S-transferase activity towards 1-chloro-2,4-dinitrobenzene in pigeon was in the order: kidney > liver > testes > brain > lung> heart. The enzyme activity with 1-chloro-2,4-dinitrobenzene as substrate was 40–44 times higher in pigeon liver and kidney than that observed with 1,2-dichloro-4-dinitrobenzene as substrate.K _m values of hepatic and renal glutathione transferase with l-chloro-2,4-dinitrobenzene as substrate were 2.5 and 3 mM respectively. Double reciprocal plots with varying reduced gluthathione concentrations resulted in biphasic curves with twoK _m values (liver 0.31 mM and 4mM; kidney 0.36 mM and 1.3 mM). The enzyme activity was inhibited by oxidized gluthathione in a dose-dependent pattern. 3-Methylcholanthrene elicited about 50% induction of hepatic glutathione transferase activity whereas phénobarbital was ineffective.     

Flight effects on certain blood parameters in homing pigeons Columba livia

• 1.1. Various blood parameters were monitored in resting and flown homing pigeons. A homing flight of 48 km lasting 60–80 min did not significantly alter plasma levels of total protein, electrolytes and plasma osmolality, which indicated maintenance of the homeostatic stability of the internal milieu during moderate exercise.
• 2.2. Plasma concentrations of marker enzymes such as alanine aminotransferase (ALAT), aspartate aminotransferase (ASAT), laetate dehydrogenase (LDH) and creatine phosphokinase (CPK) that tend to denote muscle damage and metabolic flux in prolonged exercise, were also not altered, thereby indicating the steady state of tissue structure and function during a flight of this magnitude.
• 3.3. Significant increases in plasma levels of uric acid and creatinine and decreases in plasma albumin were observed in the flown pigeons.
• 4.4. The flight-induced increase in blood uric acid could be attributed to increased purine catabolism and the increase in creatinine to increased nucleotide turnover.
• 5.5. It is suggested that the higher uric acid levels should not only enhance water conservation, but may also reduce flight-induced hyperthermia besides acting as an antioxidant defence against oxidative tissue injury.
• 6.6. The rise in creatinine is indicative of the breakdown of phosphocreatine for energy during the initial period of flight prior to the utilization of carbohydrate and lipid as fuels.
• 7.7. The decrease in plasma albumin should account for the albumin as lipid carrier lost in transport to the muscles during flight.

     

Experimental life cycle of Ascaridia columbae in intravenously infected pigeons, Columba livia.

The fate and routes of migration of infective larvae of Ascaridia columbae were studied in intravenously (i.v.) infected pigeons. Larvae were able to complete a trachael migration, and arrived at the small intestine where they established a patent infection. Granulomas were observed and histopathologically described in the lungs of i.v. infected birds.     

Blood profile of pigeons (Columba livia) during growth and breeding

Changes in the blood profile of domestic pigeons (Columba livia) were studied during growth and breeding cycle. Counts of erythrocytes and leucocytes, and values of mean corpuscular volume (MCV), mean corpuscular haemoglobin (MCH), blood volume, plasma volume (BV), width of erythrocytes, and length, width and volume of erythrocyte nuclei of squabs almost reached adult values by the 4th week of age. During courtship and mating, while the level of plasma glucose increased, those of albumin, potassium, cholesterol, calcium and uric acid decreased. At nest-building, plasma albumin and plasma calcium increased significantly. The initial phase of incubation showed an elevation in plasma calcium and a decline in cholesterol and sodium, whereas mid-phase of incubation indicated a marked rise in cholesterol and uric acid. Terminal phase of incubation had significantly low plasma protein level. During feeding and brooding period, a significant rise in sodium, protein and glucose levels and a fall in calcium were observed. Following egg-laying, there was a significant rise in calcium and a drop in protein, haemoglobin, cholesterol, sodium and MCH values. Concomitant with the phenomenal rate of growth of squabs, their haematological indices neared adult values by the 4th week of age and during breeding activity significant changes in blood values occurred.     

The role of beta-adrenergic receptors in the cutaneous water evaporation mechanism in the heat-acclimated pigeon (Columba livia)

The effects of selective and non-selective beta-adrenergic agents on cutaneous water evaporation (CWE) were studied in hand-reared rock pigeons (Columba livia). CWE was measured by the vapor diffusive resistance method, using a transient porometer. Intramuscular and subcutaneous injections of a non-selective beta-adrenergic antagonist (propranolol) or a selective beta(2)-adrenergic antagonist (ICI-118551) to heat-acclimated (HAc) pigeons at ambient temperature (T(a)) of 24 degrees C resulted in intensive CWE. The CWE values that were triggered by propranolol and ICI-118551 (18.59+/-0.73 and 16.48+/-0.70 mg cm(-2) h(-1), respectively) were close to those induced by heat exposure (17.62+/-1.40 mg cm(-2) h(-1)). Subcutaneous administration of propranolol produced local response. Intramuscular injection of salbutamol (selective beta(2)-adrenergic agonist) to HAc pigeons drastically diminished CWE induced by either propranolol, metoprolol or heat exposure. Such manipulations also enhanced panting at relatively low T(a)s (42 degrees C). The inhibition of beta(1)-adrenergic receptors by metoprolol increased CWE, while inhibition by atenolol produced no change from basal values. This difference may be attributed to their distinctive nature in penetrating the blood-brain barrier. Our findings indicate a regulatory pathway for CWE consisting of both beta(1)- and beta(2)-adrenergic receptors. We suggest that the beta(1)-adrenergic effect is restricted mainly to the CNS, while the beta(2)-adrenergic effect takes place at the effector level. We postulate this level to be either the cutaneous microvasculature or the epidermal layer.     

The effect of alpha2-adrenergic receptors on cutaneous water evaporation in the rock pigeon (Columba livia)

The role of beta-adrenergic receptors in regulating cutaneous water evaporation (CWE) in the rock pigeon (Columba livia) is well documented. Here, we studied the involvement of the alpha2-adrenergic receptors in this cooling mechanism of the heat-acclimated (HAc) pigeon. Systemic alpha2-adrenergic activation [clonidine, 50 microg kg(-1), intramuscular (i.m.)] was found to increase CWE in heat-acclimated pigeons at an ambient temperature (T(a)) of 25 degrees C. Subcutaneous administration of the drug had no significant effect. Preinjection of an alpha2-adrenergic antagonist (yohimbine, 10 mg kg(-1), i.m.) completely prevented clonidine-induced CWE and attenuated propranolol-induced CWE by 53%. Pretreatment with a beta-adrenergic agonist (isoproterenol, 4 mg kg(-1), i.m.) abolished the effect of clonidine. None of the above treatments was found to elicit significant CWE in nonacclimated (NAc) pigeons. These findings, in addition to previously reported data, indicate a complex regulatory pathway of CWE in the heat-acclimated pigeon consisting of alpha2- and beta2-adrenergic receptors. The possible hierarchical pattern of these receptors is discussed.     

Prevalence of antibodies to Toxoplasma gondii in pigeons (Columba livia) in Taiwan

Starting from January 2004 to December 2004, 665 blood serum samples from pigeons (Columba livia) were collected from 44 pigeonaries in 11 counties and 20 regions in Taiwan. These samples were examined by latex agglutination test (LAT) for antibodies to Toxoplasma gondii by using the LAT. Antibodies were found in 4.7% (31/ 665) of pigeons at a LAT titer of 1:32 or higher. The prevalence in Taiwan was highest in the northern areas (6.0%; 13/216) and lowest in the eastern areas (1.8%; 2/111).     

In vivo strain in the humerus of pigeons (Columba livia) during flight

Longitudinal and principal strain recordings were made in vivo at three sites (dorsal, anterior, and ventral) on the humeral midshaft of pigeons executing five modes of free flight: Take-off, level flight, landing, vertical ascent, and near-vertical descent. Strains were also recorded while the birds flew carrying weights that were 33%, 50%, or 100% of their body weight. The relative distribution of strain measured at the three surface midshaft sites and across the bone's cortex was found to be similar for all flight modes. Principal strains recorded in the dorsal and ventral humerus indicated considerable torsion produced by aerodynamic loading of the wing surface posterior to the bone. Measured torsional shear strains (maximum: 2,700–4,150 μ ε during level flight) were 1.5 times greater than longitudinal strains. In addition to torsion, the humerus is also subjected to significant dorsoventral bending owing to lift forces acting on the wing during the downstroke. Analysis of the cross-sectional distribution of longitudinal strains at the humeral midshaft cortex shows that the orientation of bending shifts in a regular manner during the downstroke, indicating that the wing generates progressively more thurst (vs. lift) later in the downstroke. This shift is less during take-off and vertical ascent when greater lift is required. Peak principal and longitudinal strains increased by an average of only 50% from landing to vertical ascending flight and take-off (e.g., dorsal humerus: ?1,503 to ?2,329 μ ε) and did not exceed ?2,600 μ epsiv; at any site, even when the birds flew carrying twice their body weight. Strains recorded when birds flew at two times their body weight (100% BW load) were similar in magnitude to those recorded during vertical ascent and take-off and likely represent those developed during maximal performance. Strains developed within the midshaft were maximal in the anterodorsal and posteroventral cortices, not at the dorsal, ventral, and anterior sites at which strain was recorded. Consequently, maximum strains experienced by the bone are probably 20–25% greater than those recorded (ca. 3,200 μ ε), indicating a safety factor of about 3.5 for compressive strain failure. The much higher shear strains, however, indicate a lower safety factor (1.9), in which the bone's torsional strength is its most critical design feature. Finally, the magnitude and distribution of strains developed in the humerus of pigeons are generally similar to those recorded in the humerus of large fruit-eating bats during flight. © 1995 Wiley-Liss, Inc.     

Total body haematocrit iron kinetics and erythrocyte life span in pigeons (Columba livia)

1. The mean pigeon erythrocyte life span was found to be 17-25 days by Cr51-labeled erythrocytes and 21 +/- 3.4 days by iron kinetics. 2. Total red blood cell volume has been calculated by Cr51-labeled erythrocytes while total plasma volume was determined both by a dye method and iron kinetic data. From these results total blood volume and total body haematocrit were found to be 0.090 +/- 0.002 ml/g body wt and 36 +/- 4.3%, respectively. 3. Venous haematocrit, haemoglobin concentration, erythrocyte count, mean corpuscular haemoglobin concentration, plasma iron and red blood cell iron have also been measured. 4. A significant difference between total body and venous haematocrit and a short mean red blood cell life span, due to ageing and to random destruction of erythrocytes were shown. 5. The above observations are compared with analogous available data for human beings and their physiological significance is discussed.     

Alteration in haematocrit values and plasma protein fractions during the breeding cycle of female pigeons, Columba livia

Haematocrit measurements and plasma protein electrophoresis were carried out in female pigeons (Columba livia) during the breeding cycle. Non-laying birds showed higher (p < 0.001) haematocrit values than those involved in courtship and mating, nest-building, incubation and feeding and brooding. Significant differences in relative percentage of various plasma protein fractions were observed as a function of breeding. Haematocrit could be a useful index to the non-laying status of pigeons like that in chickens, Gallus domesticus.     

Absolute and relational control of a sequential auditory discrimination by pigeons (Columba livia)

Recent evidence indicates that pigeons can readily learn visual discriminations based on both absolute and relational stimulus factors. To examine how these two types of control function in their non-dominant auditory modality, we tested four pigeons in a go/no-go sequential auditory discrimination in which both absolute and relational cues were redundantly available. In this task, sequences of different sounds created from one set of pitches were reinforced, while different sequences created from another set of pitches and any same sequences made from either set of pitches were not. Across three experiments, we independently varied the relative discriminability of the absolute and relational components. The pigeons were consistently and primarily controlled by the absolute fundamental pitch of our notes in all of the experiments, although this was influenced by the range and arrangement of the pitches used in each set. A majority of the pigeons also demonstrated relational control when this component was made more salient. The more robust control exhibited by absolute factors is consistent with the comparative hypothesis that birds in general may have a well-developed aptitude for processing absolute pitch in many auditory settings. The relational control is consistent with our recent evidence of same/different auditory learning by pigeons.     

Role of dopamine in the modulation of adrenal gland responses in the osmotically stressed pigeons, Columba livia

Salt loading on pigeons (C. livia) had stimulatory effects on brain amines (dopamine and 5-hydroxytryptamine), corticosterone, norepinephrine and epinephrine contents of adrenal gland. Conjoint administration of dopamine with hypertonic saline restored the brain amines and corticosterone of adrenal gland, but had no effect on catecholamine (CAM) contents of adrenal medulla. The excessive release of CAM in the plasma indicates sympathetic stimulation after both the treatments.