Assessment of lighting needs by W-36 laying hens via preference test

Light intensity, spectrum and pattern may affect laying hen behaviors and production performance. However, requirements of these lighting parameters from the hens’ standpoint are not fully understood. This study was conducted to investigate hens’ needs for light intensity and circadian rhythm using a light tunnel with five identical compartments each at a different fluorescent light intensity of <1, 5, 15, 30 or 100 lux. The hens were able to move freely among the respective compartments. A group of four W-36 laying hens (23 to 30 weeks of age) were tested each time, and six groups or replicates were conducted. Behaviors of the hens were continuously recorded, yielding data on daily time spent, daily feed intake, daily feeding time, and eggs laid under each light intensity and daily inter-compartment movement. The results show that the hens generally spent more time in lower light intensities. Specifically, the hens spent 6.4 h (45.4%) at 5 lux, 3.0 h (22.1%) at 15 lux, 3.1 h (22.2%) at 30 lux and 1.5 h (10.3%) at 100 lux under light condition; and an accumulation of 10.0 h in darkness (<1 lux) per day. The 10-h dark period was distributed intermittently throughout the day, averaging 25.0±0.4 min per hour. This hourly light-dark rhythm differs from the typical commercial practice of providing continuous dark period for certain part of the day (e.g. 8 h at night). Distributions of daily feed intake (87.3 g/hen) among the different light conditions mirrored the trend of time spent in the respective light intensity, that is, highest at 5 lux (28.4 g/hen, 32.5% daily total) and lowest at 100 lux (5.8 g/hen, 6.7%). Hen-day egg production rate was 96.0%. Most of the eggs were laid in <1 lux (61.9% of total) which was significantly higher than under other light intensities (P<0.05). Findings from this study offer insights into preference of fluorescent light intensity by the laying hens. Further studies to assess or verify welfare and performance responses of the hens to the preferred lighting conditions and rhythm over extended periods are recommended.     

Dopaminergic amacrine neurons of rat retinas with photoreceptor degeneration continue to respond to light

Exposure of albino rats to continuous light of low intensity (350–700 lux) for 4 months produces massive degeneration of the photoreceptor segments and cell bodies of the outer nuclear layer of the retina. Only a few heterochromatic, receptor cell nuclei remain, and no photoreceptor segments are present. On the other hand, the inner layers of these retinas remain morphologically intact. The inner nuclear layer of the normal rat retina contains a group of amacrine cells which contain the putative neurotransmitter, dopamine (DA). Short term exposure to light (30 or 60 min) markedly stimulates the rate of DA turnover in these cells in normal, previously dark-adapted rats. Such enhancement of the rate of neurotransmitter turnover in the brain has been correlated with an increase in nerve impulse activity. The present study was undertaken to determine if the dopaminergic amacrine cells of the inner nuclear layer were still responsive to light in the retinas of rats whose photoreceptors were previously destroyed by long term exposure to continuous illumination. One week before sacrifice, the animals which had been housed in continuous light for 4 months were returned to normal 14 hr light: 10 hr dark lighting conditions. At the end of this time they and a group of control rats which had been housed in cyclic lighting conditions for the entire 4 months were dark adapted for approximately 15 hr. Then the rate of retinal DA turnover was estimated from the depletion of DA following inhibition of DA synthesis by α methyl para-tyrosine. The turnover of DA in the dark-adapted retinas of the control rats and of experimental rats with photoreceptor degeneration was dramatically enhanced 2–4 fold by short term exposure (up to 1 hr) to light. Since rats are nocturnal and avoid light, we tested the light aversion of another group of rats which had been exposed to light for 4 months and then returned to cyclic lighting conditions for one week. These rats and control animals which had been maintained in cyclic lighting conditions for 4 months both chose the dark side of a light-dark box over 80% of the time. This behavior of the rats with retinal degeneration was taken as a crude indication of their continued ability to detect light. The light-induced increase in DA activity in retinas with photoreceptor degeneration may play a role in the continued ability of these rats to perceive light.     

Effects of continuous light and darkness on the eyes of the troglobitic salamander Typhlotriton spelaeus

Larval Typhlotriton spelaeus collected from five caves in Pulaski Co., Missouri, were kept as larvae or induced to transform in darkness or continuous fluorescent illumination. Larvae maintained in darkness for 215 and 279 days had smaller eyes, smaller rod inner and outer segments, and fewer metaphase figures in the genninative zone of the neural retina than comparable larvae maintained in light (258 lux). Except for visual cell size, differences were small and for each characteristic exceptions were observed. One larva kept in light showed early retinal degeneration comparable to that in transformed adults of T. spelaeus. All larvae exhibited optomotor behavior both before and after the experiment. Among animals induced to transform by L-thyroxin and maintained in darkness 111 to 366 days, visual cell and pigment epithelium degeneration was more extensive and more frequent than in animals kept for the same length of time in light (237-298 lux). In darkness the frequency of animals with retinal degeneration increased between 111 and 366 days. In light some animals exhibited pigment epithelium reduction with normal visual cells, and others had free, pigmented cells in the subretinal space. These effects were not comparable to degeneration in darkness. Eyelids covered the eyes of only a few animals in both light and dark treatments. The extent of eyelid encroachment over the eye was greater in darkness than in light. Most animals exhibited optomotor responses after experiments, but responses of animals kept in darkness were impaired in comparison to those of animals kept in light.     

Dark requirement for cell regeneration and colony formation by mesophyll protoplasts ofNicotiana plumbaginifolia Viv.

Summary The protoplasts ofNicotiana plumbaginifolia required darkness for cell regeneration and colony formation. Maximal plating efficiency of the protoplasts could be achieved by keeping the cultures in dark instead of light or dark/light sequence. Only two days of darkness prior to the illumination at 400 or 3,000 lux resulted in appreciable plating efficiency, than those of light from the beginning, but these values could not match the high plating efficiency in total darkness.     

Persistence of the Potassium Uptake Rhythm in the Presence of Exogenous Sucrose in Lemna gibba G3

The potassium uptake rhythm in a flow medium culture of Lemnagibba G3 persisted in darkness for 3 days, when the flow mediumcontained sucrose (1%). The rhythm was damped out after thatin darkness but it persisted longer when the plants were keptunder continuous weak light (80 lux). The rhythm was not dampedout when a daily light pulse (4,200 lux for 15 min) was applied.A single light pulse (4,200 lux for 15 min) at hour 48 of theprolonged dark period caused the rhythm to start again. DCMU(1 µM) slightly reduced the amplitude of the rhythm butdid not nullify the effect of the inserted light pulse. (Received September 16, 1981; Accepted February 2, 1982)     

Effects of light and dark on photoreceptors in the polychaete annelid Nereis limnicola

Summary The effects of light and dark on photoreceptors of the brackish-water polychaete annelid Nereis Hmnicola were studied by electron microscopy. Animals dark-adapted for one or two days exhibited well-formed straight microvilli (rhabdomeres) on the sensory cell processes. Continuous illumination of worms for one or two days caused extensive breakdown of the microvilli into vesicles and debris. Thirty minutes to three h of exposure of dark-adapted animals to light produced increasing severity of degradation of photoreceptoral microvilli. Light-adapted worms placed in darkness for one-half to three h showed progressive restoration of the microvilli to the dark-adapted condition. The products of degradation were internalized by both sensory and pigmented supportive cells by phagocytosis and pinocytosis.     

The effect of light and darkness on the production of cercariae of Schistosoma haematobium from Bulinus globosus

The cercariae of Schistosoma haematobium showed a diurnal periodicity of emergence from Bulinus globosus in a twelve hour light/dark cycle. Peak emission occurred at 11.00 hrs with a smaller peak at 20.00 hrs, following the start of the period of darkness. In continuous illumination this second peak was not seen, indicating that only the morning peak is circadian in origin. The evening peak occurs in response to dark treatment and can be produced by periods of darkness ranging from eight seconds to one hour. The longer the period of dark treatment the longer the rise in output is maintained on return to light conditions. Subjection of snails to periods of dark treatment during the normal light period caused a reduction in the evening peak with the largest effect seen following the longest period of darkness. An increased output of cercariae was seen following fifteen minutes exposure to a range of light intensities, the largest increase occurring at 10,000 and 7000 lux and complete darkness. The rapidity of this reaction to variations in light intensity suggests that the cercariae of S. haematobium are showing emergence in response to shadows.     

Active and resting states of the O2-evolving complex of photosystem II

During dark adaptation, a change in the O2-evolving complex (OEC) of spinach photosystem II (PSII) occurs that affects both the structure of the Mn site and the chemical properties of the OEC, as determined from low-temperature electron paramagnetic resonance (EPR) spectroscopy and O2 measurements. The S2-state multiline EPR signal, arising from a Mn-containing species in the OEC, exhibits different properties in long-term (4 h at 0 degrees C) and short-term (6 min at 0 degree C) dark-adapted PSII membranes or thylakoids. The optimal temperature for producing this EPR signal in long-term dark-adapted samples is 200 K compared to 170 K for short-term dark-adapted samples. However, in short-term dark-adapted samples, illumination at 170 K produces an EPR signal with a different hyperfine structure and a wider field range than does illumination at 160 K or below. In contrast, the line shape of the S2-state EPR signal produced in long-term dark-adapted samples is independent of the illumination temperature. The EPR-detected change in the Mn site of the OEC that occurs during dark adaptation is correlated with a change in O2 consumption activity of PSII or thylakoid membranes. PSII membranes and thylakoid membranes slowly consume O2 following illumination, but only when a functional OEC and excess reductant are present. We assign this slow consumption of O2 to a catalytic reduction of O2 by the OEC in the dark. The rate of O2 consumption decreases during dark adaptation; long-term dark-adapted PSII or thylakoid membranes do not consume O2 despite the presence of excess reductant.(ABSTRACT TRUNCATED AT 250 WORDS)     

Synthesis and post-translational processing of proenkephalin in light- and dark- adapted chicken retinas

Synthesis and post-translational processing of retinal proenkephalin in response to the light or darkness were studied in newly hatched chickens using chromatography, enzymatic digestion (trypsin-carboxypeptidase B) and radioimmunoassay. We found that the concentration of free [Met⁵]-enkephalin in crude retinal extracts increased with the time in the light and decreased in the dark. This effect was directly dependent on illumination, rather than the consequence of an endogenous circadian rhythm. In contrast, the total amount of cryptic [Met⁵]-enkephalin (in larger enkephalin-containing polypeptides) remained constant in all stages of the light/dark cycle. We also showed that the relative amounts of cryptic [Met⁵]-enkephalin stored at different molecular weights remained constant, and that the concentrations of three identified proenkephalin-derived peptides, [Met⁵]-enkephalin, [Leu⁵]-enkephalin and [Met⁵]-enkephalin-Arg⁶-Phe⁷, were higher in the light-adapted than in the dark-adapted retina; but the relative amounts (ratios) of the three proenkephalin-derived peptides stored in the light and in the dark were equal. These data are consistent with the hypothesis that synthesis and processing of proenkephalin proceed at a constant rate and with similar pattern independent of illumination or adaptation, whereas the processed enkephalins are released preferentially in the darkness and accumulated in the light.     

Fruiting ofLyophyllum tylicolor in plate culture on Soytoneglucose agar and urea-treated soil extract agar

Lyophyllum tylicolor, which forms mycelial basidia (and basidiospores), produced fruit-bodies when cultivated at 20°C under continuous illumination of 400–700 lux on agar plates containing Bacto-Soytone and glucose or an extract from urea-treated soil. Under these conditions, mycelial basidia were also observed on the Soytone-glucose agar, but not on the soil extract agar. In darkness, fruit-bodies and mycelial basidia were not observed on either medium. In culture on the soil extract agar, fruit-body primordia were produced at the position of the margin of the colony when it was transferred from darkness to continuous light; stipes did not elongate under illumination of ca. 2000 lux; and mycelial basidia and basidiospores, but not fruit-bodies, developed when glucose concentration in the medium was as high as 1% (w/v).     

THE LABELLING OF THE GANGLIOSIDIC FRACTION FROM BRAINS OF RATS EXPOSED TO DIFFERENT LEVELS OF STIMULATION AFTER INJECTION OF [6-3H]-GLUCOSAMINE

[6-³H] Glucosamine was injected into rats and then they were submitted for 1 h to light (about 1100 lux) and sound, while their controls were kept in darkness. Two series of experiments were done: in one, the animals received the injection intracerebrally; in the other, the injection was intraperitoneal. In both series the ratios of specific radioactivities (c.p.m./μmol of NANA) in mitochondria1 gangliosidic fractions to those in synaptosomal gangliosidic fractions were higher for rats exposed to illumination and sound ?+ 74 per cent, P < 0·01 and + 49 per cent, P < 0·001 (in respectively, experiments with intracerebral and intraperitoneal injections) than for animals kept in dark. For animals iqjected intraperitoneally the specific radioactivity of the mitochondria1 gangliosidic fraction from rats exposed to light was higher (+ 11 per cent, P < 0.05) than that obtained from animals kept in dark. For synaptosomes the specific radioactivity of the gangliosidic fraction obtained from animals exposed to light was lower (?15 per cent, P < 0.01) than that obtained from animals kept in dark. In animals kept under the experimental conditions already described for more than 1 h the differences tended to disappear. No differences were observed between the two groups of rats in the amounts of radioactivity found in the brain.     

Electrophoretic mobility of membrane fragments on a sucrose gradient. Application to isolated purple membrane fragments from Halobacterium halobium.

A simple technique for electrophoresis of particles is presented. The technique is based on running charged particles in a vertical tube along a sucrose gradient (20–50%). Purple membrane fragments from Halobacterium halobium were used to demonstrate the method. The migration of the fragments was linear with time in the region of 20 to 40% sucrose. Electrophoresis of purple membrane fragments under illumination, darkness, or darkness interrupted by short periods of illumination showed that at pH 4.5 the dark-adapted form of bacteriorhodopsin is less negative than its light-adapted form. At pH 6.5 and 8.5 no difference between these forms could be detected.     

RP2 and RPGR mutations and clinical correlations in patients with X-linked retinitis pigmentosa

We determined the mutation spectrum of the RP2 and RPGR genes in patients with X-linked retinitis pigmentosa (XLRP) and searched for correlations between categories of mutation and severity of disease. We screened 187 unrelated male patients for mutations, including 135 with a prior clinical diagnosis of XLRP, 11 with probable XLRP, 30 isolate cases suspected of having XLRP, and 11 with cone-rod degeneration. Mutation screening was performed by single-strand conformation analysis and by sequencing of all RP2 exons and RPGR exons 1-14, ORF15, and 15a. The refractive error, visual acuity, final dark-adapted threshold, visual field area, and 30-Hz cone electroretinogram (ERG) amplitude were measured in each patient. Among the 187 patients, we found 10 mutations in RP2, 2 of which are novel, and 80 mutations in RPGR, 41 of which are novel; 66% of the RPGR mutations were within ORF15. Among the 135 with a prior clinical diagnosis of XLRP, mutations in the RP2 and RPGR genes were found in 9 of 135 (6.7%) and 98 of 135 (72.6%), respectively, for a total of 79% of patients. Patients with RP2 mutations had, on average, lower visual acuity but similar visual field area, final dark-adapted threshold, and 30-Hz ERG amplitude compared with those with RPGR mutations. Among patients with RPGR mutations, those with ORF15 mutations had, on average, a significantly larger visual field area and a borderline larger ERG amplitude than did patients with RPGR mutations in exons 1-14. Among patients with ORF15 mutations, regression analyses showed that the final dark-adapted threshold became lower (i.e., closer to normal) and that the 30-Hz ERG amplitude increased as the length of the wild-type ORF15 amino acid sequence increased. Furthermore, as the length of the abnormal amino acid sequence following ORF15 frameshift mutations increased, the severity of disease increased.     

An Endogenous Rhythm in the Rate of CO2 Output of Bryophyllum: II. THE EFFECTS OF LIGHT AND DARKNESS ON THE PHASE AND PERIOD OF THE RHYTHM

The effects of light, darkness, and changes in light intensityon the phase and period of the endogenous rhythm in the rateof CO₂ output of excised leaves of Bryophyllum fedtschenkoihave been examined. The duration, intensity, and wavelength of a short light treatment,and the point in the cycle at which it is administered, determinethe degree of phase shift induced in a rhythm persisting indarkness. When light treatments of 3 and 6 hours' duration,at an intensity of 3,000 lux, are applied between the peaksthe phase is completely reset, the first post-treatment peakoccurring 18–19 hours after the end of the treatment.The degree of phase shift is therefore determined not by theduration of the treatment but by the time at which the treatmentterminates. One hour's illumination has little or no effect.The phase is unaffected when light treatments of up to 5 hours'duration at an intensity of 3,000 lux are applied at the crestof a peak. Over the range 8-3,000 lux the intensity of lightduring a 6-hour treatment applied between the peaks does notaffect the efficiency with which that treatment completely resetsthe phase. At an intensity of 2 lux, however, the phase delayis equal to the duration of the treatment. A 6-hour red-light treatment (850 ergs/cm.²/sec.) applied betweenthe peaks completely resets the phase whereas blue light (10,860ergs/cm.²/sec.) has no effect on the phase but induces a slightprotraction of the period. Moreover, continuous red light inhibitsthe rhythm, which recommences in blue light. A rhythm is induced in illuminated leaves when the light intensityis either gradually or suddenly reduced by at least 80 per cent.Whether a given intensity of illumination inhibits or permitsthe persistence of a rhythm depends upon the light intensityby which it is immediately preceded. A rhythm will persist in illuminated leaves for approximatelyas long as in leaves in darkness and the phase shows no correlationwith time of day. The period is unaffected by the intensityof white light (from 0-500 lux) to which the leaves are subjected.The duration of a short dark treatment, and the point in thecycle at which it is applied, determine the degree of phaseshift induced in a rhythm in illuminated leaves. The phase isreset when 3-, 6-, and 9-hour dark treatments are applied atthe crest of a peak, the amount of phase shift increasing toa maximum with 9 hours' darkness. The phase shift is not equalto the duration of the treatment. The phase is unaffected when3- and 6-hour dark treatments are applied between the peaks. The variation in the sensitivity of the phase of a rhythm persistingin darkness to short light treatments is in the opposite senseto that of a rhythm persisting in light to short dark treatments.The phase of a rhythm in illuminated leaves is completely resetwhen the leaves are transferred to continuous darkness commencingeither at the crest of, or between, the peaks. The results are discussed and compared with those of other authors.     

强光照和全黑暗条件下荒漠沙蜥视网膜内GAP-43表达的免疫组织化学

采用免疫组织化学技术研究了在强光照和全黑暗条件下荒漠沙蜥(Phrynocephalus prezewalskic)视网膜内生长相关蛋白GAP-43的表达变化。结果表明,在正常光照条件下,视网膜内GAP-43阳性表达部位主要存在于内网层;强光照条件下,GAP-43免疫染色部位主要出现在内网层、节细胞层和内核层的部分细胞核。在全黑暗条件下,在视纤维层和内网层呈阳性染色;提示视网膜在不同环境条件下GAP-43的不同定位,可能与其在相应的环境下参与不同的视觉功能有关。     

Germination characteristics ofDiamorpha cymosa seeds and an ecological interpretation

Summary Laboratory-stored seeds ofDiamorpha cymosa (Nutt.) Britton (Crassulaceae) were germinated at monthly intervals starting shortly after maturity in late May and ending at approximately the time germination is completed in the field (November). Seeds were placed at 5, 10, 15, 20, 25, 30, 15/6, 20/10, 30/15 and 35/20°C at a 14-hr photoperiod (12/12 hr thermoperiods at the alternating temperature regimes) and in constant darkness. In June, seeds were almost completely dormant and thus germinated poorly or not at all under all conditions. As seeds aged from late May to November 1. germination at the 14-hr photoperiod increased in rate and total percentage, 2. the maximum germination temperature increased from 15 to 25°C at constant temperatures and from 20/10 to 30/15°C at the alternating temperature regimes and 3. the optimum temperature for germination increased from 15 to 15–20°C at constant temperatures but remained at20/10°C at alternating temperature regimes throughout the study. During the same period germination in constant darkness was negligible at constant and alternating temperature regimes. This pattern of physiological after-ripening apparently is an adaptation to summer-dry,winter-wet habitats such as rock outcrops of southeastern United States.A short period of illumination with white light given after a 12-hr imbibition period in darkness promoted germination in the dark at 25/10°C but not at 15 or 25°C. A short period of illumination given during the imbibition period was much less effective in promoting germination in the dark. Drying up to 7 days did not cause light-stimulated seeds to lose their ability to germinate in darkness. The light requirement for seed germination probably does not play a role in restrictingD. cymosa to its well-lighted habitats on granite and sandstone outcrops.This research was supported by funds from the University of Kentucky Research Foundation and by an NIH Biomedical Sciences Support Grant to the University of Kentucky.     

Effects of changes in environmental lighting upon levels of hydroxyindole O-methyltransferase activity in the developing-chick gland.

The level of hydroxyindole O-methyltransferase (HIOMT) activity in the pienal gland of developing chicks raised under constant illumination rose more rapidly and to higher values than in the gland of birds maintained in constant darkness. Rates of net increase in activity, and levels of activity attained, for birds raised under a diurnal cycle of illumination were intermediate between those maintained in constant light or darkness. Under each of the lighting conditions, the course of increase in enzymic activity was markedly affected by variations in an unidentified factor, the source of which appeared to be the hatching eggs. Birds transferred from constant light to the dark showed either an arrest of increase in enzyme activity or a loss of activity until the levels equalled that observed for chicks of the same age raised in constant darkness. Chicks transferred from constant darkness to constant illumination showed marked increases in levels of enzyme activity at rates comparable with the maximal values observed with birds maintained under constant illumination, regardless of age and without delay. No diurnal cycle in level of HIOMT activity was observed in the pineals of 15-day birds.     

Effect of light periodicity and intensity on the growth and survival of Heterobranchus longifilis Val. 1840 (Teleostei: Clariidae) larvae after 14 days of rearing

Heterobranchus longifilis Val. 1840 larvae were reared under two light intensities, 30 lux and 915 lux, and at varying photoperiods. Results show that maximum survival (82.5 ± 6.5% respectively) at 30 lux was obtained at continuous illumination [24 h light (L)], while the minimum (65 ± 21.2%) was at the 6 h L : 18 h dark (D) treatment. Survival at 24 h D averaged 71.3 ± 6.3%, with no significant difference (P < 0.05) in growth of larvae. Maximum larval survival at 915 lux was 87.5 ± 17.7% at the 18 h L treatment. Growth was not significantly different (P < 0.05) in the treatments. Comparison of the two light intensities showed that survival was better at a photoperiod above 12 h irrespective of intensity, while growth was significantly better at the 915 lux intensity.     

Light-induced increase in the number and activity of ribosomes bound to pea chloroplast thylakoids in vivo

Within 8 to 10 minutes of illumination, chloroplast thylakoids of pea (Pisum sativum) became enriched 30 to 100% in ribosomes bound by nascent chains. Following (or, in some experiments, coincident with) this apprarent redistribution was a 25 to 65% increase in the total bound ribosome population, which was then maintained at this higher level during the normal light period. On transfer of plants to darkness, the bound ribosome population decreased to the lower dark level. White, blue (400 to 520 nanometers), and orange (545 to 690 nanometers) light were all effective in producing an increase in the bound ribosome population. The level of bound ribosomes in the oldest leaves of 16-day-old plants was 15-fold less than in the still-maturing leaf but was still increased by illumination.     

The effects of moderately high temperature on zeaxanthin accumulation and decay

Moderately high temperature reduces photosynthetic capacities of leaves with large effects on thylakoid reactions of photosynthesis, including xanthophyll conversion in the lipid phase of the thylakoid membrane. In previous studies, we have found that leaf temperature of 40°C increased zeaxanthin accumulation in dark-adapted, intact tobacco leaves following a brief illumination, but did not change the amount of zeaxanthin in light-adatped leaves. To investigate heat effects on zeaxanthin accumulation and decay, zeaxanthin level was monitored optically in dark-adapted, intact tobacco and Arabidopsis thaliana leaves at either 23 or 40°C under 45-min illumination. Heated leaves had more zeaxanthin following 3-min light but had less or comparable amounts of zeaxanthin by the end of 45?min of illumination. Zeaxanthin accumulated faster at light initiation and decayed faster upon darkening in leaves at 40°C than leaves at 23°C, indicating that heat increased the activities of both violaxanthin de-epoxidase (VDE) and zeaxanthin epoxidase (ZE). In addition, our optical measurement demonstrated in vivo that weak light enhances zeaxanthin decay relative to darkness in intact leaves of tobacco and Arabidopsis, confirming previous observations in isolated spinach chloroplasts. However, the maximum rate of decay is similar for weak light and darkness, and we used the maximum rate of decay following darkness as a measure of the rate of ZE during steady-state light. A simulation indicated that high temperature should cause a large shift in the pH dependence of the amount of zeaxanthin in leaves because of differential effects on VDE and ZE. This allows for the reduction in ΔpH caused by heat to be offset by increased VDE activity relative to ZE.