The retinal pigment epithelium of the teleost Notopterus notopterus (Pallas): appearance of basal infoldings during prolonged dark-adaptation

In teleosts, the basal part of the retinal pigment epithelium (RPE) is relatively smooth, i. e., it is free of basal membrane infoldings. In the featherback, Notopterus notopterus, whereas this is the situation in light adaptation, during dark-adaptation, especially when kept for prolonged periods (6-9 hour), numerous infoldings appear at the basal region, as found uniquely by transmission electron microscopy. In this teleost, during retinomotor movements, the rods move vitreally during dark-adaptation, while the cones do not elongate, and remain stationary in both light- and dark-adaptation. The significance of the appearance of basal infoldings in dark-adapted RPE is explained in terms of the pattern of retinomotor responses and the features of RPE and photoreceptors in this species. It is suggested that (1) the thick, impervious tapetal layer present in the RPE, (2) the unusual position of the photoreceptors in the visual cell layer of dark-adapted retina, and (3) the presumably high demand for glucose and O2 of the outer retina during dark-adaptation might contribute to cause this phenomenon in this species. The available evidence tend to associate this phenomenon with the involvement of the RPE in nutrient and O2 delivery to the photoreceptors via the basal infoldings of the RPE in dark-adapted state in this species. This has not been reported for any other teleosts to date.     

Fine structure of the retina of black bass, Micropterus salmoides (Centrarchidae, Teleostei).

The structure of light- and dark-adapted retina of the black bass, Micropterus salmoides has been studied by light and electron microscopy. This retina lacks blood vessels at all levels. The optic fiber layer is divided into fascicles by the processes of Müller cells and the ganglion cell layer is represented by a single row of voluminous cells. The inner nuclear layer consists of two layers of horizontal cells and bipolar, amacrine and interplexiform cells. In the outer plexiform layer we observed the synaptic terminals of photoreceptor cells, rod spherules and cone pedicles and terminal processes of bipolar and horizontal cells. The spherules have a single synaptic ribbon and the pedicles possess multiple synaptic ribbons. Morphologically, we have identified three types of photoreceptors: rods, single cones and equal double cones which undergo retinomotor movements in response to changes in light conditions. The cones are arranged in a square mosaic whereas the rods are dispersed between the cones.     

Daily torpor in free-ranging rock elephant shrews, Elephantulus myurus: a year-long study

Under laboratory conditions, rock elephant shrews, Elephantulus myurus, use daily torpor under both short and long photoperiod acclimation. However, use of heterothermy often differs under field and laboratory conditions. We investigated the use of torpor in free-ranging elephant shrews from May 2001 to May 2002. The elephant shrews were capable of daily torpor throughout the year, with torpor most prevalent during winter. We recorded two torpor bouts during early summer (November). We recorded a total of 467 torpor bouts during the year. The mean torpor minimum body temperature (Tbmin) for the whole year was 15.3 degrees +/-4.4 degrees C, and the mean bout length was 8.6+/-3.5 h. These values were in the range expected for daily heterotherms. However, there was some marginal overlap with hibernation characteristics; a few torpor bouts were longer than 24 h in duration, and Tbmin decreased below 10 degrees C. Torpor was highly correlated with low ambient temperature and photoperiod. Torpor was also correlated with invertebrate abundance after controlling for photoperiod effects. During the year in which this study was conducted, the rainfall was 14% below long-term average. Historical rainfall records show that summer rainfall during strong El Nino years is up to 40% below the long-term average. During these drought years, the frequency of summer torpor may be higher, highlighting the need for long-term physiological data in free-ranging animals.     

Increased air temperature during simulated autumn conditions does not increase photosynthetic carbon gain but affects the dissipation of excess energy in seedlings of the evergreen conifer Jack pine

Temperature and daylength act as environmental signals that determine the length of the growing season in boreal evergreen conifers. Climate change might affect the seasonal development of these trees, as they will experience naturally decreasing daylength during autumn, while at the same time warmer air temperature will maintain photosynthesis and respiration. We characterized the down-regulation of photosynthetic gas exchange and the mechanisms involved in the dissipation of energy in Jack pine (Pinus banksiana) in controlled environments during a simulated summer-autumn transition under natural conditions and conditions with altered air temperature and photoperiod. Using a factorial design, we dissected the effects of daylength and temperature. Control plants were grown at either warm summer conditions with 16-h photoperiod and 22 degrees C or conditions representing a cool autumn with 8 h/7 degrees C. To assess the impact of photoperiod and temperature on photosynthesis and energy dissipation, plants were also grown under either cold summer (16-h photoperiod/7 degrees C) or warm autumn conditions (8-h photoperiod/22 degrees C). Photosynthetic gas exchange was affected by both daylength and temperature. Assimilation and respiration rates under warm autumn conditions were only about one-half of the summer values but were similar to values obtained for cold summer and natural autumn treatments. In contrast, photosynthetic efficiency was largely determined by temperature but not by daylength. Plants of different treatments followed different strategies for dissipating excess energy. Whereas in the warm summer treatment safe dissipation of excess energy was facilitated via zeaxanthin, in all other treatments dissipation of excess energy was facilitated predominantly via increased aggregation of the light-harvesting complex of photosystem II. These differences were accompanied by a lower deepoxidation state and larger amounts of beta-carotene in the warm autumn treatment as well as by changes in the abundance of thylakoid membrane proteins compared to the summer condition. We conclude that photoperiod control of dormancy in Jack pine appears to negate any potential for an increased carbon gain associated with higher temperatures during the autumn season.     

Daily and seasonal activity patterns of the muskrat Ondatra zibethicus as revealed by radiotelemetry

Activity patterns of the muskrat Ondatra zibethicus L. were examined during summer and winter in Delta Marsh, Manitoba, Canada (50°11'N, 98°23'W). Animals exhibited intermittent activity throughout the day with a mean periodicity close to 6 h in both seasons. The 24-h activity pattern in summer was typically bimodal with major peaks occurring between sunset and sunrise. Muskrats were more diurnal in winter when maximal levels of activity occurred in late afternoon and early evening (1500–2000). Pronounced inter- and intra-individual variability in daily activity ensured continuous, or nearly continuous occupation of winter shelters by a variable number of muskrats, and thus enchanced microclimate stability. Daily variation in activity and weather were weakly correlated both summer and winter. Of the environmental variables tested, only photoperiod and wind speed in summer, and photoperiod and air temperature in winter correlated significantly with total time spent in lodge or burrow per day.     

Circadian rhythms in the green sunfish retina

We investigated the occurrence of circadian rhythms in retinomotor movements and retinal sensitivity in the green sunfish, Lepomis cyanellus. When green sunfish were kept in constant darkness, cone photoreceptors exhibited circadian retinomotor movements; rod photoreceptors and retinal pigment epithelium (RPE) pigment granules did not. Cones elongated during subjective night and contracted during subjective day. These results corroborate those of Burnside and Ackland (1984. Investigative Ophthalmology and Visual Science. 25:539-545). Electroretinograms (ERGs) recorded in constant darkness in response to dim flashes (lambda = 640 nm) exhibited a greater amplitude during subjective night than during subjective day. The nighttime increase in the ERG amplitude corresponded to a 3-10-fold increase in retinal sensitivity. The rhythmic changes in the ERG amplitude continued in constant darkness with a period of approximately 24 h, which indicates that the rhythm is generated by a circadian oscillator. The spectral sensitivity of the ERG recorded in constant darkness suggests that cones contribute to retinal responses during both day and night. Thus, the elongation of cone myoids during the night does not abolish the response of the cones. To examine the role of retinal efferents in generating retinal circadian rhythms, we cut the optic nerve. This procedure did not abolish the rhythms of retinomotor movement or of the ERG amplitude, but it did reduce the magnitude of the nighttime phases of both rhythms. Our results suggest that more than one endogenous oscillator regulates the retinal circadian rhythms in green sunfish. Circadian signals controlling the rhythms may be either generated within the eye or transferred to the eye via a humoral pathway.     

Circadian clock of Drosophila montana is adapted to high variation in summer day lengths and temperatures prevailing at high latitudes

Photoperiodic regulation of the circadian rhythms in insect locomotor activity has been studied in several species, but seasonal entrainment of these rhythms is still poorly understood. We have traced the entrainment of activity rhythm of northern Drosophila montana flies in a climate chamber mimicking the photoperiods and day and night temperatures that the flies encounter in northern Finland during the summer. The experiment was started by transferring freshly emerged females into the chamber in early and late summer conditions to obtain both non-diapausing and diapausing females for the studies. The locomotor activity of the females and daily changes in the expression levels of two core circadian clock genes, timeless and period, in their heads were measured at different times of summer. The study revealed several features in fly rhythmicity that are likely to help the flies to cope with high variation in the day length and temperature typical to northern summers. First, both the non-diapausing and the diapausing females showed evening activity, which decreased towards the short day length as observed in the autumn in nature. Second, timeless and period genes showed concordant daily oscillations and seasonal shifts in their expression level in both types of females. Contrary to Drosophila melanogaster, oscillation profiles of these genes were similar to each other in all conditions, including the extremely long days in early summer and the cool temperatures in late summer, and their peak expression levels were not locked to lights-off transition in any photoperiod. Third, the diapausing females were less active than the non-diapausing ones, in spite of their younger age. Overall, the study showed that D. montana clock functions well under long day conditions, and that both the photoperiod and the daily temperature cycles are important zeitgebers for seasonal changes in the circadian rhythm of this species.     

Seasonal changes in the thermoregulation of laboratory golden hamsters during acclimation to seminatural outdoor conditions

Proper adjustments of the thermoregulatory mechanisms ensure survival in the natural environment. In the present study, we tested the hypothesis that laboratory golden hamsters (Mesocricetus auratus) housed under seminatural outdoor conditions are able to acclimatize to daily and seasonal changes in the environment despite their long history of breeding in captivity. The animals experienced natural changes in the photoperiod and ambient temperature characteristic for central Poland. During experiments in the thermal gradient system, the daily rhythms of body temperature (measured as the temperature of brown adipose tissue, TBAT), preferred ambient temperature (PTa) and activity were measured in summer, autumn and spring. We found that mean TBAT was highest in autumn and least in summer, reflecting seasonal changes in the capacity for nonshivering thermogenesis (NST). In summer, TBAT followed the robust daily rhythm with the amplitude of 1.1+/-0.1 degrees C. This amplitude was depressed in autumn (0.2+/-0.1 degrees C) and partially restored in spring (0.4+/-0.1 degrees C). Seasonal changes in the daily amplitude of TBAT recorded during both transitional periods, i.e., in autumn and spring, seem to be associated with hamsters' hibernation. In autumn, mean daily PTa was lower than in summer and spring, indicating the lowering of a set point for core body temperature (Tb) regulation. Locomotor activity was much higher in spring than in summer and autumn, and it always predominated at night. We conclude that laboratory golden hamsters housed under seminatural conditions express daily and seasonal changes in the thermoregulatory mechanisms that, despite long history of breeding in captivity, enable proper acclimatization to seasonally changing environment and ensure successful hibernation and winter survival.     

Features of the daily rhythms of blood melatonin and glucose in goats during different natural photoperiod

The scope of the present study was to explore the features of the daily rhythms of plasma melatonin and blood glucose in goats during different natural photoperiod imposed by the four seasons. Five Sarda breed pluriparus female goats, 2 years old, with a mean body weight 40.5 ± 2.3 kg, lead to a natural pasture, were cannulated the day before the start of the study for sample collection performed during spring and autumn equinoxes, and winter and summer solstices for a 48-h period. On blood samples, melatonin and glucose concentrations were assessed. Our results indicate a daily rhythm of both parameters studied which characteristics were influenced by the different photoperiod due to season. During the experimental period, the two parameters showed a negative correlation, despite the onset of melatonin concentration was linked to the sunset, whereas the onset and offset of glucose were not linked to the photoperiod. In conclusion, we can claim that the two parameters are driven by different entrainable systems, but further studies are necessary to establish their correlation.     

Dopaminergic Regulation of Cone Retinomotor Movement in Isolated Teleost Retinas: I. Induction of Cone Contraction Is Mediated by D2 Receptors

In the retinas of lower vertebrates, retinal photoreceptors and melanin pigment granules of the retinal pigment epithelium (RPE) undergo characteristic movements in response to changes in light intensity and to signals from an endogenous circadian clock. To identify agents responsible for mediating light and/or circadian regulation of these retinomotor movements, we investigated the effects of hormones and neurotransmitters on cone, rod, and RPE movements in the green sunfish, Lepomis cyanellus. We report here that 3,4-dihydroxyphenylethylamine (dopamine) mimics the effect of light by inducing light-adaptive retinomotor movements in all three cell types. In isolated dark-cultured retinas, dopamine induced light-adaptive cone contraction with a half-maximal effect at 10(-8) M. This effect of dopamine was inhibited by antagonists with a potency order characteristic of D2 receptor mediation. The dopamine uptake blocker benztropine also induced light-adaptive cone contraction in isolated dark-cultured retinas, suggesting that there is continuous dopamine release in the dark but that concomitant uptake normally prevents activation of cone contraction. That dopamine plays a role in light regulation of cone movement is further suggested by the observation that light-induced cone contraction was partially inhibited by sulpiride, a selective D2 dopamine antagonist, or by Co2+, a blocker of synaptic transmission. Sulpiride also promoted dark-adaptive cone elongation in isolated light-adapted retinas, suggesting that continuous dopamine action is required in the light to maintain the light-adapted cone position. Dopamine can act directly on D2 receptors located on rod and cone inner/outer segments: dopamine induced light-adaptive retinomotor movements in isolated distal fragments of dark-adapted photoreceptors cultured in the dark. Together our results indicate that dopamine induces light-adaptive retinomotor movements in cones, rods, and RPE cells by activating D2 receptors. We suggest that, in vivo, dopamine plays a role in both light and circadian regulation of retinomotor movements.     

Retinal photoreceptor fine structure in the red-backed salamander (Plethodon cinereus).

The retinal photoreceptors of the red-backed salamander (Plethodon cinerus) have been studied by light and electron microscopy. Rods and single cones are present in this duplex retina in a ratio of about 25:1. The photoreceptors in this amphibian species are much larger than is reported for most vertebrates. In the light-adapted state, rods reach deep into the retinal epithelial (RPE) layer. The rod outer segment is composed of discs of uniform diameter displaying several very deep incisors. The rod inner segment displays a distal elliposid of mitochondria and a short stout myoid region. Rod nuclei are electron dense and often protrude through the external limiting membrane. Rod synaptic spherules are large and display several invaginated synaptic sites as well as superficial synapses. It is felt that the rods do not undergo retinomotor movements. The cone photoreceptors are much smaller than the rods and display a tapering outer segment, an unusual modified ellipsoid and a large parabolid of glycogen in the inner segment. Cone nuclei are less electron dense than rods and are located at all levels within the outer nuclear layer. The synaptic pedicle of the cones is larger, more electron lucent and display more synaptic sites (both invaginated and superficial) than that of rods. It is felt that cone photomechanical responses are minimal.     

Preliminary investigations of the daily and seasonal locomotor activity rhythms of the spiny lobster,Panulirus argus

1. Group activity rhythms of Panulirus argus in the laboratory, as measured by constant recording actographs, are described under natural photoperiod and constant temperature for a seven‐month period. Adjunct behavioral observations are reported.

2. The data indicate seasonally varying daily rhythms are present in Panulirus argus. Starting times and duration of activity periods follow a seasonal trend ‐ the animals becoming active earlier, and maintaining their activity longer, during the spring and early summer months.

3. It is suggested that the evening's high level of activity is associated with increased social interaction and that photoperiod is an important factor in the daily and seasonal components Of the rhythm.     

Effect of Sub-Polar Gyre, North Atlantic Oscillation and ambient temperature on size and abundance in the Icelandic Arctic fox

Animals are exposed to environmental factors that influence their life history and body size. Here we used the Arctic fox ( Vulpes lagopus ) as an indicator of the complex links between largescale environmental variables that influence both marine and tundra trophic dynamics to demonstrate how they affect the fox's body size and abundance. The Arctic fox inhabits throughout Iceland, where it preys mainly on birds. We studied the effects of the Sub-Polar Gyre (SPG), winter and summer North Atlantic Oscillation (NAO), mean annual winter and summer temperature, and geographic sector (eastern and western Iceland, which differ in their ecology) on variations in mandible size (6345 specimens) and body mass (2732 specimens) as well as abundance on the Arctic fox in Iceland. We found that (a) SPG index negatively affected male mandible length as well as body mass of both sexes. SPG was also negatively related to fox abundance. (b) Summer NAO had a negative effect on Arctic foxes, that is, cold summers were correlated with shorter mandibles and lower body mass. (c) Winter NAO had a significant negative effect (although weaker than that of summer NAO) on female mandible length, but not on body mass. (d) Summer temperature had a positive effect on female mandible length, but no effect on body mass. However, winter temperature had no effect on either the mandible or body mass. (e) Foxes in the eastern sector had shorter mandibles and were of lighter mass than those in the western sector. We suggest that climate conditions during the growth period of the young affected their final size both directly, by influencing energy metabolism for maintenance, but mainly through their effects on food availability. As far as we are aware, this is the first report that the SPG has an effect on vertebrates, let alone terrestrial ones.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Factors affecting plasticity in whole-organism thermal tolerance in common killifish (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>)

We characterized the degree of plasticity in thermal tolerance (assessed as critical thermal maxima; CTMax) and the relationship between thermal tolerance and underlying physiological and biochemical factors in two subspecies of a teleost fish, Fundulus heteroclitus. CTMax was not affected by repeated daily heat shock, but increased within a few days in response to warm acclimation. Loss of tolerance with acclimation to lowered temperatures occurred more slowly. Exposure to hypoxia decreased CTMax, and hyperoxia had no effect. CTMax showed a daily rhythm in both subspecies. Thermal acclimation changed the value of CTMax but did not affect the amplitude of the rhythm. Exposure to altered photoperiod had complex effects with a summer photoperiod producing a daily rhythm at higher CTMax than a spring photoperiod, and a winter photoperiod removing the rhythm. There was no daily rhythm in routine metabolic rate in either subspecies. There was no relationship between CTMax and the protein levels of the constitutive 70 and 90 kDa heat shock proteins (HSC70, HSP90β) in gill, or with mRNA levels of hsc70 in liver. There was a daily rhythm in the basal levels of the inducible hsp70-2 mRNA. Induction of hsp70-2 mRNA with mild heat shock occurred only in the evening and at night, and not during the day. These results demonstrate that there is substantial plasticity of thermal tolerance in killifish, and that this plasticity does not differ between subspecies. CTMax has a complex relationship with physiological and biochemical mechanisms that have been hypothesized to affect thermal tolerance.     

Mice Deficient of Glutamatergic Signaling from Intrinsically Photosensitive Retinal Ganglion Cells Exhibit Abnormal Circadian Photoentrainment

Several aspects of behavior and physiology, such as sleep and wakefulness, blood pressure, body temperature, and hormone secretion exhibit daily oscillations known as circadian rhythms. These circadian rhythms are orchestrated by an intrinsic biological clock in the suprachiasmatic nuclei (SCN) of the hypothalamus which is adjusted to the daily environmental cycles of day and night by the process of photoentrainment. In mammals, the neuronal signal for photoentrainment arises from a small subset of intrinsically photosensitive retinal ganglion cells (ipRGCs) that send a direct projection to the SCN. ipRGCs also mediate other non-image-forming (NIF) visual responses such as negative masking of locomotor activity by light, and the pupillary light reflex (PLR) via co-release of neurotransmitters glutamate and pituitary adenylate cyclase-activating peptide (PACAP) from their synaptic terminals. The relative contribution of each neurotransmitter system for the circadian photoentrainment and other NIF visual responses is still unresolved. We investigated the role of glutamatergic neurotransmission for circadian photoentrainment and NIF behaviors by selective ablation of ipRGC glutamatergic synaptic transmission in mice. Mutant mice displayed delayed re-entrainment to a 6 h phase shift (advance or delay) in the light cycle and incomplete photoentrainment in a symmetrical skeleton photoperiod regimen (1 h light pulses between 11 h dark periods). Circadian rhythmicity in constant darkness also was reduced in some mutant mice. Other NIF responses such as the PLR and negative masking responses to light were also partially attenuated. Overall, these results suggest that glutamate from ipRGCs drives circadian photoentrainment and negative masking responses to light.     

Ontogeny of the daily profile of plasma melatonin in European starlings raised under long or short photoperiods

Photoperiodic manipulation of young European starlings suggests that their reproductive physiology is incapable of responding to a short photoperiod until they are fully grown. This study aimed to determine whether the lack of response to a short photoperiod is reflected in the daily profile of plasma melatonin concentrations. Five-day-old starlings taken from nest boxes showed a significant (p < 0.0001) rhythm in plasma melatonin concentrations, with high values during night. In nestlings hand-reared from 5 days of age on a long photoperiod (LD 16:8), equivalent to natural photoperiod at the time, the amplitude of the daily rhythm in melatonin increased significantly (p < 0.01) with age until birds were fully grown (20 days old). In nestlings reared on a short photoperiod (LD 8:16), the daily melatonin profile remained almost identical to that of long photoperiod birds until they were fully grown. However, after 20 days old, the duration of elevated nighttime melatonin began to extend to encompass the entire period of darkness. In contrast, fully grown starlings transferred from a long to a short photoperiod had partially adapted to the short photoperiod after 5 days; by 10 days, the daily melatonin profile was identical to that of birds held chronically on a short photoperiod. Thus, consistent with responses of reproductive physiology, the pineal of young birds appears to be incapable of perceiving, or adapting to, a short photoperiod.     

Maternal transfer of photoperiodic information in Siberian hamsters. II. The nature of the maternal signal, time of signal transfer, and the effect of the maternal signal on peripubertal reproductive development in the absence of photoperiodic input

Peripubertal reproductive development of Siberian hamsters is controlled by maternally derived photoperiodic information and the ambient photoperiod present after weaning. Previous experiments suggested that the maternally derived information is transferred during gestation, not during lactation. Development was examined in several photoperiods following manipulation of gestational and lactational photoperiods; development was influenced by the gestational, but not lactational, photoperiod. Second, effects of the gestational photoperiod were observed in young reared in constant light (LL) from Day 15. Depriving the young of ambient photoperiodic information after Day 15 allows a more direct assessment of the signal received from their dams. Finally, melatonin injections to long-day dams, at certain times of day, caused transmission of a short-day signal to young, as evidenced by their development in LL and light-dark cycles. Thus, a maternally derived signal that is dependent on melatonin influences reproductive development of the young during gestation; the maternally directed pattern of development can subsequently be modified by the youngs' own response to ambient photoperiods after weaning.     

Annual cycles of growth rate, feeding rate, food conversion, plasma glucose and plasma lipids in a population of European sea bass (Dicentrarchus labrax L.) farmed in floating marine cages

Specific growth rate, average daily food intake, gross conversion efficiency, plasma total lipids, triglycerides, total cholesterol and glucose were evaluated annually in a population of European sea bass (Dicentrarchus labrax L.) intensively reared in floating marine cages. These data are aimed at improving productivity and allows more precise limits to be placed on the hematological parameters which are a necessary precondition for the assessment of fish condition and health. Specific growth rate (%), average daily food intake (%) and gross conversion efficiency showed relatively higher values in the warm spring and summer months. The specific growth rate correlated better with the seawater temperature and the average daily food intake than with the photoperiod. The concentrations of plasma total lipids, triglycerides and glucose were higher in summer. Plasma total cholesterol concentrations were higher during the pregametogenesis and gametogenesis periods. Plasma triglycerides correlated better with photoperiod and plasma glucose than with the seawater temperature.     

Seasonal change in xylem growth of <Emphasis Type="Italic">Pinus densiflora</Emphasis> in central Japan

This study investigated the seasonal change in xylem growth of Japanese red pine (Pinus densiflora). Wood cores were sampled at 2-week intervals from April to November in 2012 using the microcoring method. Daily increment rates of tracheid number and tree-ring width were compared with seasonal changes in daily mean temperature and photoperiod. Xylem growth started in early to late May and stopped in late October to early November. The maximum daily increment rates of tracheid number and tree-ring width were in early July. The 95 % confidence intervals of the timing of the maximum daily increment rates included the summer solstice (23 June) with the longest photoperiod, but not the warmest day (30 July). The maximum daily increment rate of xylem growth is thought to be controlled by the photoperiod rather than by temperature. The daily mean temperature exceeded 20 °C after the summer solstice, indicating that temperature is not a limiting factor for xylem growth. This study suggests that the timing of maximum daily increment rates of xylem growth of P. densiflora is controlled by the photoperiod.