Biomedical subjects

H Underwood

Publications and source records attributed to H Underwood.

At least 19 recordsLinked to original sources

PubMed2002

Pineal melatonin secretion, but not ocular melatonin secretion, is sufficient to maintain normal immune responses in Japanese quail (Coturnix coturnix japonica).

Reports that plasma melatonin is an important immune regulator in avian species have been rather sparse and contradictory. Also, the primary source of immune-modulating melatonin has yet to be determined in birds. In Japanese quail (Coturnix coturnix japonica), the pineal gland and eyes contribute roughly two thirds and one third of the melatonin found in the blood, respectively. Two experiments were conducted to evaluate melatonin as an immune modulator in Japanese quail and to determine the primary source of immune-modulating melatonin in this species. Experiment 1 was designed to evaluate the involvement of the pineal gland and the eyes in immunocompetence. Each of three groups of quail was assigned a surgical treatment and the cellular and humoral immune responses were determined 8 weeks following surgery. The surgical treatments were pinealectomy (Px), sham pinealectomy (SH-Px), and ocular enucleation (eye removal (Ex)). Experiment 2 utilized exogenous melatonin as a replacement to reconstitute immune responses in surgically immunocompromised birds. In this experiment, 50.0 microg/ml of melatonin, or diluent only, was provided to Px and SH-Px birds in the drinking water ad libitum. The cellular and humoral immune responses were determined after 8 weeks of melatonin treatment. In both experiments, a cutaneous basophil hypersensitivity reaction to phytohemagglutinin was measured to evaluate the cellular immune response. To evaluate the humoral immune response, primary antibody titers were determined 7 days postintravenous injection with a Chukar red blood cell suspension. Flow cytometric analysis of peripheral blood lymphocytes was performed to determine the relative percentage of CD4(+) and CD8(+) T- and B-lymphocytes in all treatments of Experiment 2. In Experiment 1, both the SH-Px and Ex surgical treatments produced similar cellular and humoral immune responses, and these responses were significantly greater than those in Px-treated birds. Pinealectomy significantly reduced the cellular and humoral immune responses from SH-Px by 25.8% and 41.3%, respectively. In Experiment 2, Px again resulted in depressed cellular and humoral immune responses. In addition, Px significantly reduced CD8(+) T-lymphocyte numbers compared to SH-Px, while B-lymphocytes remained unchanged. Melatonin administration to Px birds increased the cellular (32.9%) and humoral (30.6%) immune responses to the level of control (SH-Px) birds, although this reconstitution was not due to increased CD8(+) T- or B-lymphocytes. From these data, it was clear that removal of the pineal gland, but not the eyes, reduced cellular and humoral immune responses, which were reconstituted to normal levels by exogenous melatonin. These data suggest that immunodepression is only observed in birds with two thirds of the plasma melatonin removed by pinealectomy. Removal of one third of the plasma melatonin (by ocular enucleation) is not sufficient to reduce cellular and humoral responses in the Japanese quail.

Animals↗

PubMed2001

Circadian organization and the role of the pineal in birds.

All organisms exhibit significant daily rhythms in a myriad of functions from molecular levels to the level of the whole organism. Significantly, most of these rhythms will persist under constant conditions, showing that they are driven by an internal circadian clock. In birds the circadian system is composed of several interacting sites, each of which may contain a circadian clock. These sites include the pineal organ, the suprachiasmatic nucleus (SCN) of the hypothalamus, and, in some species, the eyes. Light is the most powerful entraining stimulus for circadian rhythms and, in birds, light can affect the system via three different pathways: the eyes, the pineal, and extraretinal photoreceptors located in the deep brain. Circadian pacemakers in the pineal and in the eyes of some avian species communicate with the hypothalamic pacemakers via the rhythmic synthesis and release of the hormone melatonin. Often the hypothalamic pacemakers are unable to sustain persistent rhythmicity in constant conditions in the absence of periodic melatonin input from the pineal (or eyes). It has also been proposed that pineal pacemakers may be unable to sustain rhythmicity in constant conditions without periodic neural input from the SCN. Significant variation can occur among birds in the relative roles that the pineal, the SCN, and the eyes play within the circadian system; for example, in the house sparrow pacemakers in the pineal play the predominant role, in the pigeon circadian pacemakers in both the pineal and eyes play a significant role, and in Japanese quail ocular pacemakers play the predominant role.

Animals↗

PubMed2000

Circadian ovulatory rhythms in Japanese quail: role of ocular and extraocular pacemakers.

Previous studies have shown that the circadian system of Japanese quail is composed of multiple photic inputs and multiple oscillators. Among these are extraretinal photoreceptors that mediate both circadian and photoperiodic responses and circadian pacemakers in the eyes that, via neural and hormonal outputs, help to maintain rhythmicity of central circadian clocks (presumably located in the suprachiasmatic area of the hypothalamus). Furthermore, a component of the central circadian system is influenced by reproductive hormones. Under certain conditions, the circadian system of female quail can be induced to split into two circadian components: one driven by ocular pacemakers and one driven by feedback from reproductive hormones. Importantly, ovulation is either inhibited or permitted as these two oscillators (or sets of oscillators) constantly change internal phase relationships with each other, suggesting an "internal coincidence" mechanism in the control of ovulation. The oviposition patterns of quail in light-dark (LD) cycles also support an internal coincidence mechanism. The authors tested the hypothesis that the ocular pacemakers are an important component of an internal coincidence mechanism controlling ovulation by examinig the effects of blinding by complete eye removal (EX), and the effects of eye-patching, on the body temperature and oviposition patterns of quail exposed to 24-h LD cycles. They also examined the effects of EX on quail exposed to continuous light (LL) and to continuous darkness (DD). Neither EX nor eye-patching affected the oviposition patterns of birds in LD. Furthermore, robust body temperature and oviposition rhythms continued in EX birds in LL, but body temperature became arrhythmic in DD with the cessation of ovulation. The results do not show a role for ocular pacemakers in the control of ovulation, but they do support the hypotheses that (1) entrainment of the central oscillators by extraretinally perceived light is sufficient to preserve a normal ovulatory pattern in LD in the absence of the ocular pacemakers, and (2) in LL, feedback of reproductive hormones onto the central oscillators is sufficient to organize the circadian system even in the absence of the ocular pacemakers. Whether or not the ocular pacemakers are normally involved in the control of ovulation is still an open question.

Animals↗

PubMed1999

Dopamine mediates circadian rhythms of rod-cone dominance in the Japanese quail retina.

A circadian clock modulates the functional organization of the Japanese quail retina. Under conditions of constant darkness, rods dominate electroretinogram (ERG) b-wave responses at night, and cones dominate them during the day, yielding a circadian rhythm in retinal sensitivity and rod-cone dominance. The activity of tyrosine hydroxylase, the rate-limiting enzyme in dopamine synthesis, also exhibits a circadian rhythm in the retina with approximately threefold higher levels during the day than at night. The rhythm of tyrosine hydroxylase activity is opposite in phase to the circadian activity of tryptophan hydroxylase, the first enzyme in the melatonin biosynthetic pathway. We tested whether dopamine may be related to the physiological rhythms of the retina by examining the actions of pharmacological agents that effect dopamine receptors. We found that blocking dopamine D2 receptors in the retina during the day mimics the nighttime state by increasing the amplitude of the b-wave and shifting the retina to rod dominance. Conversely, activating D2 receptors at night mimics the daytime state by decreasing the amplitude of the b-wave and shifting the retina to cone dominance. A selective antagonist for D1 dopamine receptors has no effect on retinal sensitivity or rod-cone dominance. Reducing retinal dopamine partially abolishes rhythms in sensitivity and yields a rod-dominated retina regardless of the time of day. These results suggest that dopamine, under the control of a circadian oscillator, has a key role in modulating sensitivity and rod-cone dominance in the Japanese quail retina.

Adaptation, Ocular↗

PubMed1999

Effects of fasting on the circadian body temperature rhythm of Japanese quail.

The effect of food deprivation on the body temperature and activity rhythms of quail was assessed in birds exposed to both light-dark (LD) cycles and to continuous darkness (DD). Quail normally exhibit a daily rhythm of body temperature in LD that will persist in DD (that is, the rhythm is circadian). In LD, 3 days' food deprivation caused the body temperature to drop below its normal nighttime levels, whereas daytime body temperature was unaffected. In DD, food deprivation caused the body temperature to drop below normal at all phases of the circadian rhythm of body temperature. Accordingly, the lack of hypothermia during the light phase of the LD cycle following food deprivation must represent a direct exogenous or "masking" effect of light, and is not an endogenous property of the circadian system. Blind birds exposed to LD 12:12 exhibited an entrained body temperature rhythm, and food deprivation caused a drop in body temperature below normal levels during both the light and dark phases of the LD cycle. Accordingly, the masking effects of light observed in normal birds on LD cycles is mediated via retinal photoreceptors and not via extraretinal photoreceptors. Measurements of activity levels before and during fasting indicate that fasting-induced hypothermia cannot be explained simply as a consequence of decreases in activity levels. Food deprivation was also observed to cause significant phase shifts in the endogenous rhythm of body temperature.

Animals↗

PubMed1999

Formal properties of the circadian and photoperiodic systems of Japanese quail: phase response curve and effects of T-cycles.

A role for the circadian system in photoperiodic time measurement in Japanese quail is controversial. The authors undertook studies of the circadian and photoperiodic system of Japanese quail to try to identify a role for the circadian system in photoperiodic time measurement. The circadian studies showed that the circadian system acts like a low-amplitude oscillator: It is readily reset by light without significant transients, has a Type 0 phase response curve (PRC), and has a large range of entrainment. In fact, a cycle length that is often used in resonance protocols (LD 6:30) is within the range of entrainment. The authors employed T-cycle experiments; that is, LD cycles with 6- and 14-h photoperiods and period lengths ranging from 18 to 36 h to test for circadian involvement in photoperiodic time measurement. The results did not give evidence for circadian involvement in photoperiodic time measurement: T-cycles utilizing 6-h photoperiods were uniformly noninductive (that is, did not stimulate the reproductive system), whereas T-cycles utilizing 14-h photoperiods were inductive (stimulatory). A good match was observed between the phase-angles exhibited on the T-cycles employing 6-h photoperiods and the predicted phase-angles calculated from a PRC generated from 6-h light pulses.

Animals↗

PubMed1997

Eye and gonad: role in the dual-oscillator circadian system of female Japanese quail.

Experiments were conducted to determine the anatomic and physiological basis of the dual-oscillator circadian system of female Japanese quail. After blocking of ocular light perception by eye-patching, the circadian body temperature rhythm dissociates into two circadian components in continuous lighting (LL). One component free runs with a period significantly shorter than 24 h [mean period (tau) = 22.7 h] and is driven by an ocular pacemaker, whereas the other component free runs with a period significantly longer than 24 h (tau = 26.3 h). The long free-running rhythm is driven by the same circadian clock that drives the circadian rhythm of ovulation. The expression of the long free-running rhythm in LL depends on the presence of the ovary: body temperature rhythmicity is abolished by ovariectomy. The two free-running oscillators in eye-patched birds showed evidence of mutual interaction. Significantly, the phase relationships that occur as the two oscillators interact can determine whether or not ovulation occurs. The results are discussed in terms of an "internal coincidence" mechanism for photoperiodic time measurement.

Animals↗

PubMed1995

Daily melatonin infusions entrain the locomotor activity of pinealectomized lizards.

Previously, it was shown that the locomotor activity rhythms of pineal-intact lizards (Sceloporus occidentalis) could be entrained to a periodicity of 24 h by 10-micrograms melatonin injections administered every other day at the same time. The present study examined the response of the circadian activity rhythm of pinealectomized S. occidentalis to daily 12-h infusions of smaller quantities of melatonin (0.1 or 5 micrograms melatonin/day). The results show that entrainment is achieved by infusion of 0.1 microgram of melatonin/day in pinealectomized lizards, as well as by 5 micrograms of melatonin/day in pinealectomized and pineal-intact lizards. Serum melatonin levels in pinealectomized lizards receiving 0.1 microgram melatonin/day (measured in the middle of the infusion period) were comparable to mid-dark levels in intact lizards. These results provide further support for the hypothesis that the pineal, via its daily rhythm of melatonin secretion, plays an important role in the circadian organization of lizards.

Animals↗

PubMed1995

The circadian rhythm of thermoregulation in Japanese quail. II. Multioscillator control.

Most biochemical, physiological, and behavioral processes in vertebrates show significant daily rhythms. Under constant conditions, these rhythms exhibit an endogenous periodicity around 24 h showing that they are driven by an internal circadian clock. In Japanese quail, the circadian clock driving activity and body temperature rhythms is functionally organized as a dual-oscillator system. Under certain conditions, such as switching birds from light:dark (LD) 12:12 to continuous darkness (DD), the body temperature rhythm splits into two circadian components that free-run independently before recoupling in a normal phase-relationship. The behavior of the activity rhythm parallels that of the body temperature rhythm, supporting the hypothesis that both rhythms are driven by the same set of oscillators. In some instances, recoupling fails to occur and birds continue to exhibit two circadian components that free-run independently. Dual-oscillator control of body temperature was observed in normal birds, pinealectomized birds, and optic nerve sectioned birds. However, birds were rendered arrhythmic by complete eye removal. It is proposed that the central circadian system (suprachiasmatic nuclei?) acts as a complex pacemaker that is functionally organized as two sets of oscillators and that circadian input from the eyes is necessary to preserve the integrity of this complex pacemaker.

Animals↗

PubMed1995

The circadian rhythm of thermoregulation in Japanese quail: III. Effects of melatonin administration.

Recent studies indicate that the circadian pacemakers in the eyes of Japanese quail are coupled to the rest of the circadian system by both neural and hormonal outputs. The effects of exogenous melatonin administration on circadian body temperature and activity rhythms of quail were tested to determine whether melatonin could be the hormonal link involved. Continuous melatonin administration caused arrhythmicity or period changes in the body temperature and activity rhythms of pinealectomized and sham-pinealectomized birds held in constant darkness and also significantly decreased the amplitude of the body temperature rhythms of normal birds held on light:dark 12:12. Further, melatonin entrained the body temperature and activity rhythms of normal birds when administered daily via the drinking water. The results show that melatonin can affect the circadian system of quail and support the hypothesis that melatonin is importantly involved in linking pacemakers in the eyes to the rest of the circadian system.

Animals↗

PubMed1994

The circadian rhythm of thermoregulation in Japanese quail. I. Role of the eyes and pineal.

Japanese quail exhibit a robust circadian rhythm in body temperature. This rhythm is readily entrainable by 24 h light-dark (LD) cycles and persists under constant conditions. Because both the pineal organ and the eyes have been implicated as major components of the circadian system of birds, the role of these organs in generating the rhythm of body temperature was investigated. Pinealectomy, when performed alone, had little effect on the body temperature rhythm of quail either under LD or under constant darkness (DD). Most birds subjected to optic nerve section alone remained rhythmic in DD although the "robustness" of the rhythm was decreased, and 25% became arrhythmic. Birds subjected to both pinealectomy and optic nerve section behaved similarly to birds subjected to optic nerve section alone. However, complete eye removal, when performed alone or in combination with pinealectomy, caused all birds to become arrhythmic in DD. The data support the hypothesis that the eyes are the loci of circadian pacemakers in quail that act, via both neural and hormonal outputs, to preserve the integrity of (self-sustaining or damped) circadian oscillators located elsewhere.

Animals↗

PubMed1993

Effects of nightbreak, T-cycle, and resonance lighting schedules on the pineal melatonin rhythm of the lizard Anolis carolinensis: correlations with the reproductive response.

Experiments were conducted to determine if a correlation exists between any aspect of the pineal melatonin rhythm (such as its duration or phase) in the lizard Anolis carolinensis and the reproductive response to photoperiod. The rhythm of pineal melatonin content was determined in anoles exposed to nightbreak lighting protocols (10L:5D:1L:8D, 10L:10D:1L:3D), resonance lighting cycles (LD 11:13, LD 11:25), and T-cycle lighting protocols (LD 11:7, LD 11:9, LD 11:13, LD 11:15, LD 11:19) and compared with the testicular response to these lighting protocols as determined previously [Underwood and Hyde, (1990) J. Comp. Physiol. (A) 167:231-243]. Different T-cycles and nightbreak cycles elicited changes in both the duration of the melatonin peak and the phase of the melatonin peak relative to these light cycles. The response to the resonance cycle (LD 11:25) was complex, probably due to the overlapping patterns of two groups whose pineal melatonin rhythms were entrained approximately 12 hr out of phase with each other. No correlation was observed between the duration, or the amplitude, of the nocturnal melatonin peaks seen on the various light cycles and the reproductive response to these cycles. A correlation was observed between the phase of the pineal melatonin rhythm and the reproductive response. Light cycles were inductive (stimulated testicular growth) when the entrained melatonin rhythm peaked near the light-to-dark or the dark-to-light transition, but they were not inductive when the melatonin rhythm peaked during the middle third of the night. These results suggest that if melatonin is involved in the transduction of photoperiodic information in Anolis, neither the duration nor amplitude of the nocturnal melatonin pulse is involved in the measurement of day length. Instead, the phase-relationship of the melatonin rhythm to the rest of the circadian system may determine photoperiodic responsiveness.

Animals↗

PubMed1992

The superior cervical ganglia are not necessary for entrainment or persistence of the pineal melatonin rhythm in Japanese quail.

The avian pineal exhibits a daily rhythm in the synthesis and secretion of the hormone, melatonin, which is involved in maintaining temporal order within the circadian system of some species. The pineal is richly innervated by sympathetic nerves which originate in the superior cervical ganglia (SCG) and, in the chicken, these nerves play a role in generating the melatonin rhythm. In the Japanese quail, the pineal melatonin rhythm can be entrained by light perceived directly by the pineal or by light perceived by the eyes. The role of the sympathetic innervation of the pineal was investigated in the Japanese quail by subjecting birds to bilateral superior cervical ganglionectomy (SCGX) and determining if SCGX either abolished the ability of retinally perceived light to entrain the pineal melatonin rhythm or if it disrupted the rhythm under constant darkness (DD). The results show that SCGX neither prevented entrainment of the pineal melatonin rhythm by retinally perceived light nor affected the rhythm expressed in DD. An entrainment pathway between the eyes and pineal exists in quail which does not involve the SCG.

Animals↗

PubMed1991

Retinally perceived light can entrain the pineal melatonin rhythm in Japanese quail.

The avian pineal organ contains a circadian oscillator that can drive a daily rhythm of melatonin synthesis. In some avian species the pineal organ may act, via the cyclic release of melatonin, as a pacemaker within a multioscillator circadian system. The routes by which light entrains the pineal melatonin rhythm were investigated in the Japanese quail. A 'patching' protocol was used to expose directly either the eyes or the pineal to a light-dark cycle while the rest of the bird was exposed to constant light. The results show that the pineal melatonin rhythm can be entrained (1) by light perceived directly or (2) by light perceived by the eyes. Furthermore, the pathway by which light entrains the pineal melatonin rhythm includes the optic nerves because transection of the optic nerve eliminates the ability of ocularly perceived light to entrain the pineal melatonin rhythm.

Animals↗

PubMed1990

The pineal and melatonin: regulators of circadian function in lower vertebrates.

The pineal has been identified as a major circadian pacemaker within the circadian system of a number of lower vertebrates although other pacemaking sites have been implicated as well. The rhythmic synthesis and secretion of the pineal hormone, melatonin, is suggested as the mechanism by which the pineal controls circadian oscillators located elsewhere. Both light and temperature cycles can entrain the pineal melatonin rhythm. The pineal, therefore, acts as a photo and thermoendocrine transducer which functions to synchronize internal cycle with cycles in the environment. A model is presented which portrays the pineal as a major component of a 'multioscillator' circadian system and which suggests how these multiple circadian clocks are coupled to each other and to cycles of light and temperature in the external world.

Animals↗

PubMed1990

The quail's eye: a biological clock.

The site (intraocular vs. extraocular) of the biological clock driving a rhythm in melatonin content in the eyes of Japanese quail was investigated by alternately patching the left and right eyes of individual birds, otherwise held in constant light, for 12-hr periods. This patching protocol, therefore, exposed each eye to a light-dark cycle (LD 12:12) 180 degrees (12 hr) out of phase with the LD cycle experienced by the other eye. The optic nerves to both eyes were transected prior to initiating the patching protocol. The ocular melatonin rhythm (OMR) of the left eyes of quail could be entrained by this procedure 180 degrees out of phase with the rhythm expressed by the right eyes. Since optic nerve section would have deprived any putative extraocular clocks of photic entrainment information, the results show conclusively that the clock driving the OMR is located within the eye itself. In addition, the OMR of Japanese quail is remarkably unaffected by removing two potential neural inputs to the eye (sympathetic innervation from the superior cervical ganglia, and input from the isthmo-optic nucleus of the midbrain); this suggests that these inputs are not required to maintain the OMR. Finally, the clock driving the OMR of one eye does not appear to be coupled to the clock driving the OMR in the other eye, since permanently patching one eye abolished the ability of the patched eye to re-entrain to an 8-hr shift in the phase of an LD 12:12 cycle, whereas the exposed eye rapidly re-entrained to the phase-shifted cycle.

Animals↗

PubMed1990

Melatonin does not link the eyes to the rest of the circadian system in quail: a neural pathway is involved.

Blinding by enucleation has a dramatic effect on the circadian activity rhythm of Japanese quail. The activity patterns of enucleated birds held under 24-hr light-dark cycles are disrupted, although entrainment can persist in many birds. In constant darkness (DD), blinded birds are rendered arrhythmic. These results demonstrate that the eyes are a major component of the circadian system, and that insofar as enucleation produces arrhythmicity in DD, the eyes' role is not merely a photosensory one. The eyes of quail can synthesize and secrete the hormone melatonin, which has been implicated as a blood-borne messenger relaying timing information between elements of the circadian system in some avian species. However, the way in which the eyes communicate with the rest of the circadian system in quail appears to be neural, since (1) optic nerve section produces the same effects as blinding by enucleation on the circadian activity rhythm, and (2) eyes subjected to optic nerve section retain their ability to synthesize and secrete melatonin.

Animals↗

PubMed1989

Cloning and DNA sequence analysis of a Lactococcus bacteriophage lysin gene.

A gene for the lysin of Lactococcus lactis bacteriphage phi vML3 was cloned using an Escherichia coli/bacteriophage lambda host-vector system. The gene was detected by its expression of antimicrobial activity against L. lactis cells in a bioassay. The cloned fragment was analysed by sub-cloning on to E. coli plasmid vectors and by restriction endonuclease and deletion mapping. Its entire DNA sequence was determined and an open reading frame for the lysin structural gene was identified. The sequenced lysin gene would express a protein of 187 amino acids with a molecular weight of 21,090, which is in good agreement with that of a protein detected after in vitro transcription and translation of DNA encoding the gene. Expression of the lysin gene in E. coli and B. subtilis from an adjacent bacteriophage promoter was readily detected but in L. lactis expression of lysin was found to be lethal. The bacteriophage phi vML3 lysin had sequence homology with protein 15 of B. subtilis bacteriophage PZA. This protein is involved in DNA packaging during bacteriophage maturation rather than in host cell lysis. The cloning and analysis of the phi vML3 lysin gene is of importance in further understanding lactic streptococcal bacteriophages, for the development of positive selection vectors and for biotechnological applications of relevance to the dairy industry.

Amino Acid Sequence↗