Differences In Light Sensitivity Research Articles

Dark continuous noise from visual pigment as a major mechanism underlying rod–cone difference in light sensitivity

Retinal rods and cones underlie scotopic and photopic vision, respectively. Their pigments exhibit spontaneous isomerizations (quantal noise) in darkness due to intrinsic thermal energy. This quantal noise, albeit exceedingly low in rods, dictates the light threshold for scotopic vision. The same quantal noise in cones, however, is too low to explain the much higher diurnal light threshold. Separately, a dark continuous noise is present in rods, long accepted to originate from an intrinsic random activation of the cyclic guanosine monophosphate (cGMP)-phosphodiesterase enzyme mediating phototransduction downstream of the pigment. Here, we report the surprising finding that most of this rod dark continuous noise actually originates from rhodopsin itself. Importantly, we found the same continuous noise with a much higher magnitude from cone pigments. The rod and cone continuous noises are apparently both associated with a hitherto unrecognized "metastable" pigment conformational state physiologically resembling that in apo-opsin (opsin devoid of chromophore) and is intermittently active for very brief moments. The cone holopigment's high continuous noise is expected to act as an intrinsic equivalent light and adapt the cone dramatically, accounting for a major part of the light-sensitivity difference between rods and cones in darkness.

Far-Red Light Inhibits Soybean Biomass and Yield by Modulating Plant Photosynthesis

Alterations in the light environment can significantly influence soybean morphology and yield formation; however, the effects and mechanisms of different light qualities on these aspects require further investigation. Consequently, we selected soybean cultivars with marked differences in light sensitivity as test materials, conducted experiments with red, blue, and green light qualities against a blue light background, and analyzed parameters related to leaf photosynthetic capacity, chlorophyll fluorescence, morphological characteristics, biomass, and yield variations following different light quality treatments. The results showed that following far-red light treatment, soybean plants exhibited significant shade avoidance syndrome, internode elongation, increased plant height, and a marked reduction in both root and leaf biomass, as well as total biomass. Furthermore, there was a substantial reduction in photosynthetic capacity. This indicated that far-red light exerts an inhibitory effect on soybean growth and yield formation. Red light has basically no regulatory effect on plant morphology and yield, while green light has a yield-increasing effect, but there was a cultivar effect. This study not only enhances our understanding of the mechanisms through which light quality regulates plant photosynthesis but also lays a scientific foundation for future crop light environment management and for the further exploration of light quality’s regulatory potential on crop growth.

Greater sensitivity of the circadian system of women to bright light, but not dim-to-moderate light.

Women typically sleep and wake earlier than men and have been shown to have earlier circadian timing relative to the light/dark cycle that synchronizes the clock. A potential mechanism for earlier timing in women is an altered response of the circadian system to evening light. We characterized individual-level dose-response curves for light-induced melatonin suppression using a within-subjects protocol. Fifty-six participants (29 women, 27 men; aged 18-30 years) were exposed to a range of light illuminances (10, 30, 50, 100, 200, 400, and 2000 lux) using melatonin suppression relative to a dim control (<1 lux) as a marker of light sensitivity. Women were free from hormonal contraception. To examine the potential influence of sex hormones, estradiol and progesterone was examined in women and testosterone was examined in a subset of men. Menstrual phase was monitored using self-reports and estradiol and progesterone levels. Women exhibited significantly greater melatonin suppression than men under the 400-lux and 2000-lux conditions, but not under lower light conditions (10-200 lux). Light sensitivity did not differ by menstrual phase, nor was it associated with levels of estradiol, progesterone, or testosterone, suggesting the sex differences in light sensitivity were not acutely driven by circulating levels of sex hormones. These results suggest that sex differences in circadian timing are not due to differences in the response to dim/moderate light exposures typically experienced in the evening. The finding of increased bright light sensitivity in women suggests that sex differences in circadian timing could plausibly instead be driven by a greater sensitivity to phase-advancing effects of bright morning light.

Light on Shedding: A Review of Sex and Menstrual Cycle Differences in the Physiological Effects of Light in Humans.

The human circadian system responds to light as low as 30 photopic lux. Furthermore, recent evidence shows that there are huge individual differences in light sensitivity, which may help to explain why some people are more susceptible to sleep and circadian disruption than others. The biological mechanisms underlying the differences in light sensitivity remain largely unknown. A key variable of interest in understanding these individual differences in light sensitivity is biological sex. It is possible that in humans, males and females differ in their sensitivity to light, but the evidence is inconclusive. This is in part due to the historic exclusion of women in biomedical research. Hormonal fluctuations across the menstrual cycle in women has often been cited as a confound by researchers. Attitudes, however, are changing with funding and publication agencies advocating for more inclusive research frameworks and mandating that women and minorities participate in scientific research studies. In this article, we distill the existing knowledge regarding the relationship between light and the menstrual cycle. There is some evidence of a relationship between light and the menstrual cycle, but the nature of this relationship seems dependent on the timing of the light source (sunlight, moonlight, and electric light at night). Light sensitivity may be influenced by biological sex and menstrual phase but there might not be any effect at all. To better understand the relationship between light, the circadian system, and the menstrual cycle, future research needs to be designed thoughtfully, conducted rigorously, and reported transparently.

Patches of Dysflective Cones in Eyes With No Known Disease.

PurposeTo characterize the structure and function of patches of dysflective cones in the foveal region of subjects with normal vision and no known pathology. Dysflective cones are cones that have little or no reflective properties in optical coherence tomography (OCT) or adaptive optics scanning laser ophthalmoscope (AOSLO) images yet exhibit measurable function.MethodsAOSLO images were surveyed for the presence of hyporeflective cone patches, and subjects were brought back for imaging to determine the changes in the hyporeflective region. Adaptive optics microperimetry (AOMP) was used to assess the function of hyporeflective patches in four subjects to determine that they did, in fact, contain dysflective cones. AOMP utilized a stimulus size of less than 1 arcmin to measure thresholds inside and outside the hyporeflective region.ResultsNineteen out of 47 individuals retrospectively reviewed had one or more regions with hyporeflective cone patches in one or both eyes. Ten subjects with hyporeflective cone patches were brought back for imaging. Seven of the 10 had resolved at follow up, and in three subjects new hyporeflective patches appeared in a different location. All AOMP-measured subjects had measurable function in the dysflective cone region. Three out of four subjects showed no difference in light sensitivity in the dysflective region compared to adjacent areas, and one subject showed a 3× reduction in sensitivity in the area.ConclusionsPatches of dysflective cone have been identified in subjects with normal vision and no known pathology, and we have observed instances where dysflective cones in these subjects regain normal reflective properties.

Melatonin, circadian rhythms and glaucoma: current perspective.

Melatonin, circadian rhythms and glaucoma: current perspective.

Considerations on how to light the night-shift

Electric lighting has decreased dependence on natural light to illuminate the workplace. Humans are genetically predisposed to be day-oriented (diurnal) and depend on daylight to regulate circadian rhythms. Shift work will force workers to sleep and work at non-biological times, inducing circadian disruption with implications for workers’ safety and health. The scientific literature may be used in practice in shift work settings to improve safety, performance and health in the workplace by reducing circadian misalignment. Alertness profiles at work and degree of melatonin suppression may indicate degree of circadian disruption among workers. However, when considering lighting solutions at night, there are several factors that need consideration. Light measures based on biological effectiveness should be used rather than room illuminance giving better predictions of performance and long-term health among workers. Also, large individual differences in light sensitivity and preferences suggest not only to rely on common lighting alone but also to implement complementary individual lighting solutions at work. Lighting advice should consider shift scheduling characteristics such as speed of turnover and shift timing to guide decisions of preferred circadian phase influence. Lighting should also include the flexibility to be fit for morning, afternoon and evening work.

The Role of Light Sensitivity and Intrinsic Circadian Period in Predicting Individual Circadian Timing.

There is large interindividual variability in circadian timing, which is underestimated by mathematical models of the circadian clock. Interindividual differences in timing have traditionally been modeled by changing the intrinsic circadian period, but recent findings reveal an additional potential source of variability: large interindividual differences in light sensitivity. Using an established model of the human circadian clock with real-world light recordings, we investigated whether changes in light sensitivity parameters or intrinsic circadian period could capture variability in circadian timing between and within individuals. Healthy participants (n = 12, aged 18-26 years) underwent continuous light monitoring for 3 weeks (Actiwatch Spectrum). Salivary dim-light melatonin onset (DLMO) was measured each week. Using the recorded light patterns, a sensitivity analysis for predicted DLMO times was performed, varying 3 model parameters within physiological ranges: (1) a parameter determining the steepness of the dose-response curve to light (p), (2) a parameter determining the shape of the phase-response curve to light (K), and (3) the intrinsic circadian period (tau). These parameters were then fitted to obtain optimal predictions of the three DLMO times for each individual. The sensitivity analysis showed that the range of variation in the average predicted DLMO times across participants was 0.65 h for p, 4.28 h for K, and 3.26 h for tau. The default model predicted the DLMO times with a mean absolute error of 1.02 h, whereas fitting all 3 parameters reduced the mean absolute error to 0.28 h. Fitting the parameters independently, we found mean absolute errors of 0.83 h for p, 0.53 h for K, and 0.42 h for tau. Fitting p and K together reduced the mean absolute error to 0.44 h. Light sensitivity parameters captured similar variability in phase compared with intrinsic circadian period, indicating they are viable targets for individualizing circadian phase predictions. Future prospective work is needed that uses measures of light sensitivity to validate this approach.

Individual differences in light sensitivity affect sleep and circadian rhythms.

Artificial lighting is omnipresent in contemporary society with disruptive consequences for human sleep and circadian rhythms because of overexposure to light, particularly in the evening/night hours. Recent evidence shows large individual variations in circadian photosensitivity, such as melatonin suppression, due to artificial light exposure. Despite the emerging body of research indicating that the effects of light on sleep and circadian rhythms vary dramatically across individuals, recommendations for appropriate light exposure in real-life settings rarely consider such individual effects. This review addresses recently identified links among individual traits, for example, age, sex, chronotype, genetic haplotypes, and the effects of evening/night light on sleep and circadian hallmarks, based on human laboratory and field studies. Target biological mechanisms for individual differences in light sensitivity include differences occurring within the retina and downstream, such as the central circadian clock. This review also highlights that there are wide gaps of uncertainty, despite the growing awareness that individual differences shape the effects of evening/night light on sleep and circadian physiology. These include (1) why do certain individual traits differentially affect the influence of light on sleep and circadian rhythms; (2) what is the translational value of individual differences in light sensitivity in populations typically exposed to light at night, such as night shift workers; and (3) what is the magnitude of individual differences in light sensitivity in population-based studies? Collectively, the current findings provide strong support for considering individual differences when defining optimal lighting specifications, thus allowing for personalized lighting solutions that promote quality of life and health.

Pinopsin and photoreception in the pineal organ of the domestic turkey during post-hatching development

The avian pineal organ is photosensitive because of the presence of photopigments, of which pinopsin seems to be one of the most important. This organ is subject to far-reaching changes during post-hatching development, but evidence regarding pinopsin presence and direct photoreception during this time is lacking. This study was carried out to demonstrate the following: 1) the structures showing immunoreactivity to pinopsin in the turkey pineal organ, 2) the changes of these structures during development, 3) the pinopsin localization in pinealocytes in monolayer cultures, and 4) the role of direct photoreception in the regulation of melatonin secretion in pineal organs in adult turkeys. Pinopsin immunoreactivity was localized in the apical extensions of columnar cells limiting the follicular lumen, in fiber-like structures located between columnar cells in the inner part of follicle wall, in string-shapes or small spherical structures distributed in the outer part of follicle wall and in amorphous material inside the follicle lumen. In young birds, immunoreactivity was also sporadically noted in cell bodies of rudimentary receptor pinealocytes. The distribution of pinopsin showed prominent age-dependent changes, including a subsequent increase in pinopsin-positive structures in the outer part of the follicle wall and a prominent reduction in the number and size of positive apical extensions in 40- and 56-week-old turkeys. These data demonstrate that the role of secretory pinealocytes in pineal photoreception increases with age. In monolayer cultures, all pinealocytes showed strong reactions in club- or bulbous-shaped prolongations. The pineal organs of adult birds were less sensitive to light exposition at night than those of young turkeys, which points to differences in light sensitivity between rudimentary receptor and secretory pinealocytes. However, direct photoreception could play an important role in the regulation of melatonin secretion in adult turkeys.

Traits related to bipolar disorder are associated with an increased post-illumination pupil response

Mood states in bipolar disorder appear to be closely linked to changes in sleep and circadian function. It has been suggested that hypersensitivity of the circadian system to light may be a trait vulnerability for bipolar disorder. Healthy persons with emotional-behavioural traits associated with bipolar disorder also appear to exhibit problems with circadian rhythms, which may be associated with individual differences in light sensitivity. This study investigated the melanopsin-driven post-illumination pupil response (PIPR) in relation to emotional-behavioural traits associated with bipolar disorder (measured with the General Behavior Inventory) in a non-clinical group (n = 61). An increased PIPR was associated with increased bipolar disorder-related traits. Specifically, the hypomania scale of the General Behavior Inventory was associated with an increased post-blue PIPR. Further, both the full hypomania and shortened ‘7 Up’ scales were significantly predicted by PIPR, after age, sex and depressive traits were controlled. These findings suggest that increased sensitivity to light may be a risk factor for mood problems in the general population, and support the idea that hypersensitivity to light is a trait vulnerability for, rather than symptom of, bipolar disorder.

Determination of the Acceptable Ambient Light Exposure during Drug Product Manufacturing for Long-Term Stability of Monoclonal Antibodies.

Monoclonal antibodies (mAbs) are exposed to light during drug product (DP) manufacturing, and the acceptable levels of light exposure needs to be determined based on the impact on product quality. In this study, a mild and more representative light model consisting of ambient light instead of stress light as prescribed by ICH Q1B was used to evaluate the impact of light exposure on mAb DP quality. The immediate effect of ambient light exposure on protein DP quality was determined to be dependent on the amount of light exposure rather than light intensity (up to 5000 lux). The impact on quality of mAbs is product specific due to their differences in light sensitivity, in which mAb II shows larger increases in IEC basic variants and larger decreases in SEC monomer when compared to mAb I after 0.24 million lux hours of light exposure. The acceptable ambient light exposure for mAb II DP manufacturing was determined to be 0.13 million lux hours, in which no impact on product quality was observed after the short-term light exposure. Additionally, real-time storage (5 °C) of the DP after the prescribed ambient light exposure showed no impact to various product quality attributes. The light model used in this study is capable of determining the acceptable amount of ambient light exposure for mAbs, especially during DP manufacturing processes.

Sex differences in light sensitivity impact on brightness perception, vigilant attention and sleep in humans

Artificial light endows a “round-the-clock”, 24-h/7-d society. Chronic exposure to light at night contributes to health hazards for humans, including disorders of sleep. Yet the influence of inter-individual traits, such as sex-differences, on light sensitivity remains to be established. Here we investigated potential sex-differences to evening light exposure of 40 lx at 6500 K (blue-enriched) or at 2500 K (non-blue-enriched), and their impact on brightness perception, vigilant attention and sleep physiology. In contrast to women, men had higher brightness perception and faster reaction times in a sustained attention task during blue-enriched light than non-blue-enriched. After blue-enriched light exposure, men had significantly higher all-night frontal NREM sleep slow-wave activity (SWA: 2–4 Hz), than women, particularly during the beginning of the sleep episode. Furthermore, brightness perception during blue-enriched light significantly predicted men’s improved sustained attention performance and increased frontal NREM SWA. Our data indicate that, in contrast to women, men show a stronger response to blue-enriched light in the late evening even at very low light levels (40lux), as indexed by increased vigilant attention and sleep EEG hallmarks. Collectively, the data indicate that sex differences in light sensitivity might play a key role for ensuring the success of individually-targeted light interventions.

An ancestral haplotype of the human PERIOD2 gene associates with reduced sensitivity to light-induced melatonin suppression.

Humans show various responses to the environmental stimulus in individual levels as “physiological variations.” However, it has been unclear if these are caused by genetic variations. In this study, we examined the association between the physiological variation of response to light-stimulus and genetic polymorphisms. We collected physiological data from 43 subjects, including light-induced melatonin suppression, and performed haplotype analyses on the clock genes, PER2 and PER3, exhibiting geographical differentiation of allele frequencies. Among the haplotypes of PER3, no significant difference in light sensitivity was found. However, three common haplotypes of PER2 accounted for more than 96% of the chromosomes in subjects, and 1 of those 3 had a significantly low-sensitive response to light-stimulus (P < 0.05). The homozygote of the low-sensitive PER2 haplotype showed significantly lower percentages of melatonin suppression (P < 0.05), and the heterozygotes of the haplotypes varied their ratios, indicating that the physiological variation for light-sensitivity is evidently related to the PER2 polymorphism. Compared with global haplotype frequencies, the haplotype with a low-sensitive response was more frequent in Africans than in non-Africans, and came to the root in the phylogenetic tree, suggesting that the low light-sensitive haplotype is the ancestral type, whereas the other haplotypes with high sensitivity to light are the derived types. Hence, we speculate that the high light-sensitive haplotypes have spread throughout the world after the Out-of-Africa migration of modern humans.

Light-sensitive brain pathways and aging.

Notwithstanding its effects on the classical visual system allowing image formation, light acts upon several non-image-forming (NIF) functions including body temperature, hormonal secretions, sleep-wake cycle, alertness, and cognitive performance. Studies have shown that NIF functions are maximally sensitive to blue wavelengths (460–480 nm), in comparison to longer light wavelengths. Higher blue light sensitivity has been reported for melatonin suppression, pupillary constriction, vigilance, and performance improvement but also for modulation of cognitive brain functions. Studies investigating acute stimulating effects of light on brain activity during the execution of cognitive tasks have suggested that brain activations progress from subcortical regions involved in alertness, such as the thalamus, the hypothalamus, and the brainstem, before reaching cortical regions associated with the ongoing task. In the course of aging, lower blue light sensitivity of some NIF functions has been reported. Here, we first describe neural pathways underlying effects of light on NIF functions and we discuss eye and cerebral mechanisms associated with aging which may affect NIF light sensitivity. Thereafter, we report results of investigations on pupillary constriction and cognitive brain sensitivity to light in the course of aging. Whereas the impact of light on cognitive brain responses appears to decrease substantially, pupillary constriction seems to remain more intact over the lifespan. Altogether, these results demonstrate that aging research should take into account the diversity of the pathways underlying the effects of light on specific NIF functions which may explain their differences in light sensitivity.

Benchmarking of optical dimerizer systems.

Optical dimerizers are a powerful new class of optogenetic tools that allow light-inducible control of protein–protein interactions. Such tools have been useful for regulating cellular pathways and processes with high spatiotemporal resolution in live cells, and a growing number of dimerizer systems are available. As these systems have been characterized by different groups using different methods, it has been difficult for users to compare their properties. Here, we set about to systematically benchmark the properties of four optical dimerizer systems, CRY2/CIB1, TULIPs, phyB/PIF3, and phyB/PIF6. Using a yeast transcriptional assay, we find significant differences in light sensitivity and fold-activation levels between the red light regulated systems but similar responses between the CRY2/CIB and TULIP systems. Further comparison of the ability of the CRY2/CIB1 and TULIP systems to regulate a yeast MAPK signaling pathway also showed similar responses, with slightly less background activity in the dark observed with CRY2/CIB. In the process of developing this work, we also generated an improved blue-light-regulated transcriptional system using CRY2/CIB in yeast. In addition, we demonstrate successful application of the CRY2/CIB dimerizers using a membrane-tethered CRY2, which may allow for better local control of protein interactions. Taken together, this work allows for a better understanding of the capacities of these different dimerization systems and demonstrates new uses of these dimerizers to control signaling and transcription in yeast.

Male but not female Wistar rats show increased anxiety-like behaviour in response to bright light in the defensive withdrawal test

The defensive withdrawal test (DWT) is used to model anxiety-like behaviour in rats. The aim of this study was to investigate whether an aversive stimulus, bright light, affects the behaviour in this test. Additionally, the effect of habituation to the apparatus was studied. Both male and female Wistar rats were used to study whether sex differences exist in the DWT, as reported for other tests of anxiety. On day 1 half of the rats were tested under low light and half under bright light. Two to seven days after trial one the same rats were repeatedly tested under the same light condition for five consecutive days. The male rats showed a higher degree of anxiety-like behaviour when tested under bright light than under low light. In contrast, the behaviour of the female rats was not affected by changes in illumination. Male rats also exhibited elevated anxiety-like behaviour compared to female rats under bright light, whereas under low light conditions no sex difference was seen. Males in low light habituated much faster than males tested under bright light, whereas in females there was little difference in habituation between low and bright light. In summary, we found that bright light is aversive for male but not female Wistar rats in the DWT. Whether this is due to sex differences in light sensitivity or if females respond with a different behavioural strategy in response to bright light, which could not be detected in the DWT, remains to be elucidated.

Origin of the Relative Afferent Pupillary Defect in Optic Tract Lesions

To determine the percent decussation of pupil input fibers in humans and to explain the size and range of the log unit relative afferent pupillary defect (RAPD) in patients with optic tract lesions. Experimental study. Five patients with a unilateral optic tract lesion. The pupil response from light stimulation of the nasal hemifield, temporal hemifield, and full field of each eye of 5 patients with a unilateral optic tract lesion was recorded using computerized binocular infrared pupillography. Six stimulus light intensities, separated by 0.5-log unit steps, were used; 12 stimulus repetitions were given for each stimulus condition. For each stimulus condition, the pupil response of each eye was characterized by plotting the mean pupil contraction amplitude as a function of stimulus light intensity. The percentage of decussating afferent pupillomotor input fibers was calculated from the ratio of the maximal pupil contractions elicited from each eye. The RAPD was determined pupillographically from full-field stimulation to each eye. In all patients, the pupil response from the functioning temporal hemifield ipsilateral to the tract lesion was greater than that from the functioning contralateral nasal hemifield. This temporal-nasal asymmetry increased with increasing stimulus intensity and was similar in hemifield and full-field stimuli, eventually saturating at maximal light intensity. The log unit RAPD did not correlate with the estimated percentage of decussating pupil fibers, which ranged from 54% to 67%. In patients with a unilateral optic tract lesion, the pupillary responses from full-field stimulation to each eye are the same as comparing the functioning temporal field with the functioning nasal field. The percentage of decussating fibers is reflected in the ratio of the maximal pupil contraction amplitudes resulting from stimulus input between the two eyes. The RAPD that occurs in this setting reflects the difference in light sensitivity between the intact temporal and nasal hemifields. Its magnitude does not correlate with the difference in the number of crossed and uncrossed axons, but its sidedness contralateral to the side of the optic tract lesion is consistent with the greater percentage of decussating pupillomotor input.

The effect of gender on the melatonin suppression by light: a dose response relationship.

It is well known that light is an inhibitor of pineal melatonin secretion in humans. However, the effect of gender on the melatonin suppression by dim and bright light is still controversial. The present study investigated the effect of gender on the suppression of melatonin at five light intensities (0, 200, 500, 1,000, 3,000 lux). Five healthy men and women attended five testing sessions separated by one week. At each session, subjects were exposed to light from midnight to 0100 hours in a sitting position. Blood samples where collected at regular intervals and plasma melatonin concentration was measured using a specific radioimmunoassay. No gender differences were found in melatonin suppression by light at any of the five light intensities (p > 0.1). Furthermore, the mean melatonin suppression by light in both males and females was dose dependent (17% [200 lux], 40% [500 lux], 56% [1,000 lux] and 74% [3,000 lux]). Our findings suggest that melatonin suppression by light in intensity dependent, with no gender differences in light sensitivity.

Light sensitivity in larval fishes: Implications for vertical zonation in the pelagic zone

Larval fishes exhibit marked vertical zonation patterns that are important in planktonic predator‐prey interactions. This zonation has been related to both proximate physical (e.g., light intensity) and ultimate biological factors (e.g., predator risk, prey availability). We hypothesize that changes in visual sensitivity and feeding performance in larval fishes provide a constraint on depth distributions and that this constraint may represent the major determinant of vertical distribution patterns. To test the generality of the relationship between larval age and visual sensitivity, larvae were obtained from captive breeding adults of seven species (three Apogonidae and four Pomacentridae). The lowest light intensity at which feeding behavior occurred was measured using infrared video techniques with natural prey items. The visual sensitivity of the larvae, measured as initiation of feeding, increased by about 3 orders‐of‐magnitude during the larval phase. The abilities of the larvae were broadly similar within families, but there were marked differences between families. Pomacentrids were about an order‐of‐magnitude less light‐sensitive than apogonids. Larval apogonids are potentially capable of feeding at a depth of about 100 m in clear tropical waters at 4 mm standard length (SL) and to 200 m at 12 mm SL. The patterns of ontogenetic and phylogenetic differences in light sensitivity are consistent with field observations on the vertical distributions of larvae in these taxa.