text:
Differential Bird Responses to Colour Morphs of an Aposematic Leaf Beetle may Affect Variation in Morph Frequencies in Polymorphic Prey Populations
- 828 Downloads
The selection of prey by predators should, theoretically, favour uniformity in the warning signals displayed by unpalatable prey. Nevertheless, some aposematically coloured species are polymorphic. We tested the hypothesis that colour morphs of unpalatable prey differ in the efficacy of their aposematic signal for birds, thereby affecting the selective advantages of these morphs. We used colour morphs (red-and-black light, red-and-black dark and metallic) of the chemically defended leaf beetle Chrysomela lapponica. In laboratory experiments, naïve great tits (Parus major) attacked live beetles of all colour morphs at the same rate. By contrast, wild-caught tits attacked light beetles at first encounter at the same rate as a novel control prey, but significantly avoided both dark and metallic beetles. Beetles of all colour morphs were similarly unpalatable for birds, and about half of the attacked beetles were released unharmed. Avoidance learning was similarly fast for all three leaf beetle morphs. However, in the next-day memory test, the dark beetles were attacked at a greater rate than beetles of two other morphs, indicating their lower memorability. A field experiment suggests that at low C. lapponica population densities, dark beetles have a survival advantage over light beetles, potentially due to the lesser conspicuousness of the dark pattern; however, when the density is high, dark beetles lose this advantage due to the low memorability of their pattern. Thus, the direction of selective bird predation on prey colour morphs may depend on prey density and thereby contribute to temporal shifts in colour morph frequencies following population fluctuations.
KeywordsAvoidance learning Colour polymorphism Density dependence Parus major Population dynamics Selective predation
Conspicuous colouration in unpalatable or noxious animals acts as a warning signal of their unprofitability, a phenomenon known as aposematism. Both theoretical expectations (Servedio 2000; Endler and Mappes 2004) and empirical studies (Mallet and Joron 1999; Chouteau et al. 2016) point to a positive frequency-dependent (antiapostatic) selection of aposematic prey. A more common warning display is recognised more easily by the predators, so avoidance learning is faster. This results in a higher survival of prey with a more frequent form of a warning signal (Joron and Mallet 1998; Beatty et al. 2004), i.e. the fitness of a phenotype increases with its frequency. This process is responsible for the observed convergence between chemically defended aposematic species that form mimicry assemblages (Sherratt 2008; Chouteau et al. 2016), as well as for intra-specific monomorphism in warning signalling (Ruxton et al. 2004; Borer et al. 2010). However, contrary to theoretical predictions, many organisms possess variable warning signals, and intra-population variation in aposematic pattern is a frequent phenomenon in a wide range of taxa: insects, for example the hibiscus harlequin bug (Tectocoris diophthalmus; Fabricant et al. 2014), the alpine leaf beetle (Oreina gloriosa; Borer et al. 2010) and the wood tiger moth (Parasemia plantaginis; Hegna et al. 2013); amphibians, for example the dyeing poison frog (Dendrobates tinctorius; Rojas and Endler 2013); and reptiles, for example vipers (Vipera berus and V. aspis; Broennimann et al. 2014).
Several mechanisms are capable of maintaining the within-population polymorphism of colour patterns (Mallet and Joron 1999; Gordon et al. 2015). Of particular importance is negative frequency-dependent (apostatic) selection, in which predators consume disproportionally more of a particular prey when it is common and less of that prey when it is rare (reviewed by Punzalan et al. 2005). Several studies have provided evidence for apostatic selection, but most have examined repeated encounters with palatable prey. In these situations, predator develops a search image, which increases the probability of detecting that prey type (Dukas 2002; Punzalan et al. 2005). This scenario differs from the situation in which the prey advertises its toxicity or unpalatability by its conspicuous warning coloration. Encounters with unpalatable aposematic prey result in avoidance learning in the predators and lead to a positive frequency-dependent (antiapostatic) selection that affects the evolution of coloration in aposematic prey species (Borer et al. 2010; Chouteau et al. 2016).
The directions of selection on prey may change if different predators vary in their responses to a particular aposematic signal (Endler and Mappes 2004; Lindstedt et al. 2011; Nokelainen et al. 2014). The efficacy of warning displays is influenced by their conspicuousness (Endler and Mappes 2004; Guilford 1986; Gamberale-Stille 2001; Prudic et al. 2007), which depends on the context (background, illumination, distance) and pattern traits, such as colour, internal contrast, size and symmetry of the pattern elements (Forsman and Merilaita 1999; Lindstedt et al. 2008; Aronsson and Gamberale-Stille 2013). For example, some aposematic patterns are conspicuous at a close distance and become cryptic with increasing distance or against another background (Tullberg et al. 2005; Gamberale-Stille et al. 2009; Barnett et al. 2017). Colour morphs within populations of aposematic prey may differ in traits that affect their conspicuousness, and this variation may lead to interplay between positive and negative frequency-dependent selection by predators. In this way, a polymorphic prey population may escape from positive frequency-dependent selection. However, only a few studies (Lindstedt et al. 2011, 2016; Hegna et al. 2013; Nokelainen et al. 2014) have explicitly addressed the role of visually hunting predators in the maintenance of polymorphism within populations of aposematic prey. Prediction of the selective advantages of different morphs in natural populations of defended prey requires exploration of whether coexisting aposematic displays differ in their ability to elicit unlearned wariness in potential predators, accelerate the rate of avoidance learning and strengthen the memory of aversive experience with the prey.
Many leaf beetle species exhibit within-population polymorphism in their colour pattern (Mikhailov 2001; Nahrung and Allen 2005). The Lapland leaf beetle, Chrysomela lapponica, is chemically defended (Pasteels et al. 1989) and highly polymorphic in the coloration of its elytra, which varies from entirely red through various red-and-black patterns, to an entirely black or metallic colour (Milyashevich 2000; Mikhailov 2001); all these patterns can be considered as warning signals advertising the beetle’s unpalatability to predators. The intra-population polymorphism of C. lapponica differs considerably between geographic populations (Milyashevich 2000; Mikhailov 2001; Gross et al. 2004); for example, the proportion of the completely black (metallic) morph varies from zero in Fennoscandian populations (Gross et al. 2004; Zvereva et al. 2002) to 86% in the Yekaterinburg population (Y. Mikhailov, personal communication) and to 100% in the Belarusian population (Milyashevich 2000).
The existence and maintenance of dark (melanic) morphs in some populations may reflect thermoregulatory benefits of dark morphs at low temperatures (Clusella-Trullas et al. 2008). This prediction is confirmed by an increase in the proportion of melanic individuals of some species with latitude and altitude (Clusella-Trullas et al. 2008; Hegna et al. 2013) and by the concomitant decrease in the proportion of melanic morphs in populations with climate warming (Brakefield and de Jong 2011; Zeuss et al. 2014). In C. lapponica, dark males copulate more frequently than light males at low temperatures due to their superior thermal capacity, which allows them to move faster (Zverev et al. 2018). However, the proportion of black or metallic morphs in C. lapponica does not depend on the latitude or altitude of the locality: it may be high in southern low-altitude populations and low in northern and high-altitude populations (Mikhailov 2001). Thus, factors other than a thermal melanism are likely to contribute to the observed geographic variation in polymorphism of C. lapponica.
The proportions of different morphs in a population may also vary among years (Svensson and Abbott 2005; Wang et al. 2009). In particular, the proportions of dark morphs in C. lapponica populations depend on the beetle population density and stage of population cycle (Zvereva et al. 2002). These among-year variations were explained by the selective mortality of colour morphs due to density-dependent changes in plant quality (Zvereva et al. 2002), but density-dependent selective predation by visually hunting predators can also contribute to the observed variation. However, this contribution cannot be estimated yet, because the responses of avian predators to different colour morphs of C. lapponica beetles have not been studied. The extensive knowledge of the life history and population ecology of C. lapponica (Gross et al. 2004; Zvereva et al. 2002; Zverev et al. 2018) makes this species an excellent model for studies on the role of predators in the maintenance of colour polymorphism and in determining the variations in morph frequencies in aposematic prey populations.
In this study, we tested the hypothesis that avian predators respond differently to different colour morphs of aposematic prey co-existing within the same population, thereby potentially affecting the selective advantages of these morphs irrespectively of their frequencies. We studied the bird behavioural responses, namely initial wariness, avoidance learning and memory, to three colour morphs of C. lapponica and the ability of birds to generalise between the different colour morphs in the experiments with both wild-caught and hand-reared great tits, Parus major. We then linked the observed differences in great tit behaviour with the survival of different C. lapponica morphs in nature by estimating the intensity of bird attacks on leaf beetles in field experiments. We also determined whether selective bird predation depends on beetle population density by conducting field experiments in localities where this leaf beetle was common and where it was absent.
Materials and Methods
Prey and Predator
Leaf Beetle Chrysomela lapponica
Chrysomela lapponica (Chrysomelidae) is a medium-sized (5–8 mm long) leaf beetle widely distributed in Palaearctic. Geographic populations of this leaf beetle differ in their host-plant specialisation, but all populations used in this study were willow-feeding to control for potential effects of host plants on beetle chemical defences. Adults emerge in late summer, feed for a couple of weeks, hibernate in soil and start feeding and copulating on willows at the time of the next leaf flush. Larvae feed on willow foliage from early July to early August and pupate on the leaves.
Like other species of the subtribe Chrysomelina, both larvae and adults of C. lapponica are defended against natural enemies by toxic and/or deterrent chemicals, which differ in larvae and adults (Pasteels et al. 1984). Adults synthesise de novo isoxazolinone derivatives that are released from pronotal and elytral exocrine glands upon disturbance (Pasteels et al. 1989; Sugeno and Matsuda 2002) and act as a defence against predators (Pauls et al. 2016). Beetle unpalatability is advertised by two different types of aposematic coloration: a red-and-black pattern or a metallic black with blue iridescence (Fig. 1). Preliminary analysis indicates that the colour morphs of C. lapponica do not differ in the nature of their defensive chemistry (M. Hilker, personal communication).
In this study, we used beetles from three populations differing in within-population polymorphism. The population from Monchegorsk (67°58′45″N, 32°58′52″E; hereafter, the Kola population) includes patterned light and dark morphs, with the dark beetles representing 5–40% of all individuals (Zverev et al. 2018). The populations from Yekaterinburg (56°45′55″N, 60°39′19″E; Siberia population hereafter) and Medvezhjegorsk (63°00′07″N, 34°22′55″E; Karelia population hereafter) contain the typical patterned morph (corresponding to the light morphs from Kola population) and the metallic black morph; the latter accounts for ca. 80% in the Siberia population and ca. 30% in the Karelia population. Crossing experiments demonstrated that the colour patterns in C. lapponica are genetically based (Zverev et al. 2018, and unpublished).
The beetles used in our experiments were reared in the laboratory from larvae collected in July 2015 and June 2016. They were fed with their native host plants (Salix myrsinifolia for Kola and Karelia populations and S. caprea for Siberia population) until pupation. The hatched beetles were then fed with the same host plants until they discontinued feeding and were subsequently kept in vials lined with moist paper at 5 °C until the experiments. Dark and light morphs of red-and-black patterned beetles from Kola population were further distinguished based on the proportion of black coloration on the elytra (> 85% and < 70%, respectively), as described by Zverev et al. (2018); we did not use intermediately coloured individuals (about 5% of all beetles) in our experiments. EZ performed the classification of all beetles. We used live beetles in the laboratory experiments with great tits; for field experiments, the beetles were killed by freezing.
Great Tit (Parus major)
Great tits are insectivorous passerines that are widely distributed across Palaearctic over a range of woodland habitats (Cramp and Perrins 1993). Great tits live in the same habitats as C. lapponica, so they are likely to encounter these leaf beetles in nature.
Adult birds were caught using mist nets in Prague (50°04′48″N, 14°14′24″E) during September and October 2015. All wild-caught birds were at least 4 months old, and a sample contained a balanced proportion of yearlings and older individuals. The wild-caught birds were experienced with various prey types from the wild including, potentially, noxious and aposematic prey. The birds were kept separately for 2–5 days before the experiment in indoor cages (50 × 40 × 50 cm) for habituation to the laboratory environment. The cage illumination simulated daylight, and photoperiod was set according to outdoor conditions. The birds’ diet consisted of mealworms (larvae of Tenebrio molitor), sunflower seeds and a commercial food mixture (Uni patee, Orlux).
Juveniles were obtained from nest boxes placed in woods at the outskirts of Prague in spring 2016. We took birds at the age of 12–15 days, when they were naïve with respect to any kind of unpalatable prey. Birds were hand-reared until they were able to feed independently. Their diet consisted of mealworms and a commercial mixture for hand-rearing the passerines (Handmix, Orlux). Nestlings were kept in artificial nest boxes in small groups. After fledging, they were housed and fed in the same way as adult birds. Experiments with juveniles started after the 35th day of their life, when they were fully independent.
Facilities Used in Experiments with Caged Birds
Experiments were conducted at the Faculty of Science, Charles University, Prague. The experiments with wild-caught birds were conducted during September–October 2015, and experiments with naïve, hand-reared birds during July 2016. We tested the birds individually in wooden cages (70 × 70 × 70 cm) with wire mesh walls, equipped with a perch, a dish with water and a rotating feeding tray. The cage was illuminated by a daylight-simulating Biolux Combi 18-W bulb (Osram). The tested prey was offered to the birds in a glass Petri dish (50 mm in diameter) placed on the rotating tray. Before the experiment, the birds were habituated to the experimental cages, were trained to take food from a glass Petri dish placed on a rotating tray, and were deprived of food for 2 h. Prey items were offered on a green background made from a leaf (white beam Sorbus aria in 2015, goat willow S. caprea in 2016). We observed the birds through a one way glass in the front wall of the cage. We recorded all experiments with a video camera and continuously noted the behavioural elements using Observer XT 8.0 (Noldus).
Experiments with Wild-Caught Adult Great Tits
The birds (63 in total) were divided into three experimental groups: 20 birds were tested with beetles of light morph, 22 birds with dark morph and 21 birds with metallic morph. We used the nymphs of Jamaican field crickets (Gryllus assimilis) as a control palatable prey that would be unfamiliar to the birds to account for the effects of prey novelty on bird responses.
An avoidance-learning session consisted of a series of trials conducted immediately one after another. We defined a trial as offering our target prey (i.e. a beetle of a particular colour morph) to the bird. Each trial was preceded by offering a control prey with the aim of controlling the bird’s foraging motivation. The trial was terminated after the bird consumed the prey; otherwise, it lasted 5 min.
If the bird did not attack the beetle during the first trial (first-offer trial hereafter), we continued the trials (i.e. offered another cricket and another beetle) until the beetle was attacked. If the bird did not attack the beetle after 20 trials, this bird was excluded from the avoidance-learning session and, subsequently, from the memory test.
The trial in which the bird attacked the beetle was recorded as the first trial of the avoidance-learning session. After this trial, we continued offering crickets and beetles, in turn, with up to three trials with each prey.
The day after the avoidance-learning session, we checked whether the birds remembered their experience with a particular prey. This memory test consisted of one trial: after the bird consumed a cricket, it was offered a beetle of the same morph as had been offered on the previous day.
The generalisation test was carried out immediately after the memory test. After the bird consumed a cricket, we simultaneously offered the bird two beetles of two different morphs that had not been used in the avoidance-learning session and the memory test. The beetles were placed in two separate Petri dishes on a rotating tray, and the trial lasted a maximum of 10 min.
During each trial we recorded: (1) the number of approaches to prey and time spent for close prey inspection before making the decision about the attack or retreat; (2) the time from the beginning of the first-offer trial to the first attack on prey (first-attack latency hereafter); (3) whether prey items were attacked (touched, pecked or seized), killed and eaten by birds; and (4) the duration of discomfort behaviour (cleaning the bill, shaking the head, ruffling the feathers, vomiting). If, at the end of a trial, the beetle was damaged by a bird but still alive, we recorded whether it survived until the next day. If the beetle died during this time, we considered it as ‘killed’.
Experiment with Naïve Juvenile Great Tits
The birds (44 in total) were divided into two experimental groups: 21 birds were tested with beetles of light and dark morphs, and 23 birds with light and metallic morphs. These combinations of morphs reflect the situation that occurs in natural populations with two kinds of polymorphism, in which either the metallic and the light-patterned morphs (Siberia and Karelia populations) or the dark-patterned and the light-patterned (Kola population) morphs coexist.
Experiments were conducted in the same way as the generalisation test, i.e. we used a dual-choice test. We first offered a mealworm (a familiar control palatable prey), then a cricket (a novel control palatable prey). If the bird ate the cricket, we offered it two C. lapponica beetles, each belonging to a different morph, and recorded which morph was attacked first. We also recorded the same behavioural elements as in the experiment with wild-caught birds. This trial lasted 10 min.
Two field experiments were conducted to test whether wild birds cause different mortality in beetles with different colour patterns and whether selective predation depends on the novelty of this type of prey, which decreases with an increase in prey population density. Morphs occurring in the Kola population were tested at four sites in the Kola Peninsula. The study area is inhabited by C. lapponica, but its distribution is patchy and the selected sites differ considerably in the abundance of this leaf beetle (Zvereva et al. 2002). The population density of C. lapponica in two our sites (67°36′N, 32°39′E and 67°35′N, 32°33′E) was extremely low: no beetles were found during a 30-min survey on host plants in June of 2016 (low density sites hereafter). In two other sites (67°48′N, 32°47′E and 67°38′N, 32°45′E) C. lapponica was more abundant: 6 and 8 beetles were found during a 30-min survey (high density sites hereafter). Five S. myrsinifolia bushes were selected at each site, and beetles from Kola population, killed by freezing and freshly defrosted, were glued with silicone to the leaves (four light and three dark beetles per plant, one beetle per leaf). The experiment ran from 11 to 29 August 2016.
Morphs occurring in the Karelia population were tested in the surroundings of Turku, at sites where C. lapponica has never been recorded and thus was a novel prey for local birds. We selected 10 bushes of S. caprea and attached beetles from the Karelia population in the same way as described for the Kola Peninsula (seven light patterned and three metallic beetles per plant). The experiment ran from 21 July to 2 September 2016.
In both experiments, we checked beetles at 3–5 days intervals and individually recorded the damage caused by birds as follows: 0, no damage; 1, beetle is decapitated, or the elytra or thorax bear visible damage; 2, both elytra or whole beetle is missing; 3, signs of two attacks are recorded during the observation time.
The numbers of birds that attacked, killed or ate the prey, as well as the numbers of beetles that survived bird’s attack, were compared by Fisher’s exact test (2 × 2 contingency tables) or by the Chi square test (3 × 2 contingency tables). Attack latencies were compared between beetles and control prey by the Wilcoxon signed rank test (paired differences); unpaired differences in attack latencies between beetle colour morphs were analysed by the Wilcoxon rank sum test. The durations of different behavioural elements were compared using the Kruskal–Wallis test. In field experiments, tree-specific mean scores were calculated for each morph and compared using a paired t-test. All reported P values are two-tailed; the calculations were performed in R 3.0.2 and SAS 9.4.
Reactions of Wild-Caught Adult Birds to Colour Morphs of C. lapponica
First Reaction to Prey
Birds that attempted an attack hesitated longer before attacking dark (Wilcoxon signed rank test: V = 30, P = 0.004) and metallic (V = 3, P < 0.001) beetles than before attacking crickets, i.e. novel control prey (Fig. 2). In contrast, first-attack latencies did not differ between beetles of the light morph and crickets (V = 105, P = 1). First-attack latencies differed between the beetle colour morphs: birds hesitated less before attacking light beetles than before attacking dark (Wilcoxon rank sum test: W = 87, P = 0.0018) and metallic (W = 60, P = 0.0021) beetles (Fig. 2).
Beetles of the light morph were attacked in the first-offer trial more frequently compared to beetles of dark and metallic morphs (Fig. 3a; Fisher’s exact test: P = 0.0007 and P = 0.0001, respectively).
The numbers of birds that did not attack any beetle during 20 subsequent trials were similar for light and dark beetles (0 and 2 birds, respectively; Fisher’s exact test: P = 0.26), and greater for metallic than for light morphs (6 and 0 birds, respectively; P = 0.01). Across all groups, 45.5% of the birds attacked a beetle only in one trial. After the first attack, the attack rates significantly decreased for all colour morphs (Fisher’s exact test, light morph: P = 0.003; dark morph: P = 0.01; metallic morph: P = 0.04). The proportion of birds that attacked the beetles in more than one trial did not differ among the morphs (frequency test: χ2 = 2.82, df = 2, P = 0.24; Fig. 3b). The same numbers of birds killed and consumed beetles of all colour morphs during the avoidance-learning session (Supplementary material Table 1). The numbers of beetles that survived bird attack did not differ among colour morphs (light morph: 41.0%; dark morph: 37.8%; metallic morph: 56.5%; frequency test: χ2 = 1.48, df = 2, P = 0.48).
The number of bird approaches to the prey differed depending on the beetle colour morph (Kruskal–Wallis test: χ2 = 7.07, df = 2, P = 0.03), as did the duration of close prey inspection by the bird before attack or retreat (χ2 = 12.2, df = 2, P = 0.002), which was longer for dark morphs than for either light (χ2 = 5.76, df = 1, P = 0.02) or metallic morphs (χ2 = 11.2, df = 1, P = 0.0008) (Supplementary material Table 2). Nevertheless, the number of attacks per bird did not differ between the beetle morphs (Supplementary material Table 2; χ2 = 1.73, df = 2, P = 0.42). The same was true for the proportions of ‘negative’ decisions—when the birds rejected the prey after approaching it and inspecting it from a short distance (χ2 = 2.08, df = 2, P = 0.35). Across all trials, the birds hesitated longer before approaching (χ2 = 4.91, df = 1, P = 0.03) and attacking (χ2 = 4.07, df = 1, P = 0.04) the metallic beetles when compared to the patterned beetles (light and dark morphs combined) (Supplementary material Table 2).
After attacking a beetle, the birds spent more time cleaning their bills when compared to birds that did not attack a beetle (χ2 = 23.4, df = 1, P < 0.0001; Supplementary material Table 2). Only slight symptoms indicating toxic effects were observed after beetle consumption and only in three birds; this behaviour consisted of sitting briefly on a perch with their feathers ruffled, after which the birds started to behave normally again.
The frequencies of the bird attacks in the memory test (Fig. 3b; frequency test: χ2 = 7.61, df = 2, P = 0.022) and the numbers of attacks per trial (Kruskal–Wallis test: χ2 = 8.48, df = 2, P = 0.01) differed among the beetle morphs (Supplementary material Table 1). Fewer birds attacked beetles of the light and metallic morphs in the memory test than they did in the first-offer trial (Fisher’s exact test, light morph: P < 0.001; metallic morph: P = 0.002; Fig. 3a, b). By contrast, birds did not appear to remember their experience with the dark morph, as the attack rates were similar for the first-offer trial and for the memory test (P = 0.21, Fig. 3a, b). Consequently, in the memory test, the birds attacked the dark morph more frequently than the light and metallic morphs (P = 0.003; Fig. 3b).
Birds that learned to avoid the metallic morph attacked fewer beetles altogether in the generalisation test than did the birds that learned to avoid one of the patterned morphs (Fisher’s exact test, metallic vs dark beetles: P = 0.015; metallic vs light beetles: P = 0.026; Fig. 4). Birds that previously attacked patterned morphs (either light or dark) attacked the metallic morph more frequently than did birds that had previous experience with the metallic morph itself (P = 0.04). This effect was not symmetrical: birds that had experience with the metallic morph subsequently avoided both patterned morphs at the same rate as they avoided the metallic morph (P = 1; Fig. 4).
Reactions of Naïve Juvenile Birds to Colour Morphs of C. lapponica
The attack latencies did not differ between novel and familiar control prey (crickets and mealworms, respectively; Wilcoxon signed rank test: V = 115, P = 0.11; Fig. 5), indicating a low degree of neophobia. Moreover, the attack latencies did not differ between beetles and crickets either in birds choosing between dark and light beetles (V = 35.5, P = 0.23) or in those choosing between metallic and light beetles (V = 56, P = 0.07) (Fig. 5). The first attack latencies did not differ either between light and dark morphs (Wilcoxon rank sum test: W = 13.5, P = 0.65) or between light and metallic morphs (W = 26, P = 0.44) (Fig. 5).
All naïve birds attacked at least one of the two offered beetles; therefore, the attack rate was significantly higher when compared with that of the wild-caught birds tested in a similar dual-choice test following their experience with C. lapponica (effect of experience, Fisher’s exact test: P < 0.0001).
In total, 92% of offered beetles were attacked by juvenile birds. Neither the first choice (Fisher’s exact test: P = 0.84 and P = 0.51 for birds selecting metallic vs light morphs and dark vs light morphs, respectively) nor beetle mortality (P = 1 and P = 0.57, respectively) depended on the colour morph of the beetles (Supplementary material Table 3). The proportion of attacked beetles that were subsequently killed by the birds did not differ between the colour morphs (P = 0.75 and P = 0.20 for birds selecting metallic vs light morphs and dark vs light morphs, respectively; Supplementary material Table 3). However, the birds tested with metallic vs light morphs approached the light beetles more frequently than they approached the metallic ones (Wilcoxon signed rank test: V = 33, P = 0.03); the number of bird approaches per trial did not differ between the light and dark morphs (V = 22.5, P = 0.30) (Supplementary material Table 3).
Across all morphs, 44.4% of the beetles survived the attack by juvenile birds, and this value did not differ from the survival of beetles attacked by adult birds (43.4%; P = 1). The percentage of beetles that were at least partly consumed by birds was also similar for both juvenile and adult birds (38.3 and 28.3%, respectively; P = 0.19) across all experiments.
Twenty-six of 44 birds demonstrated discomfort behaviour (mostly cleaning the bill) after attacking a beetle, but none of juvenile birds that consumed the beetle showed any symptoms of intoxication.
Bird Predation on Colour Morphs of C. lapponica in Field Experiments
In the Kola Peninsula, birds damaged similar numbers of beetles in habitats with low and high densities of C. lapponica (34.3 and 30.0%; Fisher’s exact test: P = 0.72). However, birds attacked beetles more intensively in sites with low density of C. lapponica when compared with high density sites (Kruskal–Wallis test: χ2 = 4.36, df = 1, P = 0.037; Fig. 6). In low density sites, the attacks were more intensive on the light morphs than on the dark morphs (paired test: t = 2.54, df = 7, P = 0.037; Fig. 6), whereas in high density sites, the attack rates were similar for both morphs (t = 0.66, df = 5, P = 0.56; Fig. 6).
In Turku, where C. lapponica did not occur, 48% of all beetles were damaged; and light beetles were damaged more frequently than metallic beetles (t = 2.82, df = 8, P = 0.023; Fig. 6).
Chemical Defence of C. lapponica Against Bird Predators
In our experiments, almost half of the attacked beetles were released unharmed, indicating that beetle chemical defence enhances individual survival upon predator attack. Moreover, only ca. 30% of the attacked beetles were eaten by great tits, confirming that these birds find C. lapponica beetles unpalatable. Thus, the chemical defence of C. lapponica appeared to be mostly based on taste rejection, and on irritating rather than on toxic effects, as indicated by the repeated bill cleaning by the birds after contact with beetle. Actual visible effects of poisoning were rare and weak. This mechanism of defence against birds is likely to be typical for many leaf beetle species of subtribe Chrysomelina, which share the same defensive chemicals (Pasteels et al. 1984).
Our findings support the conclusion by Skelhorn and Rowe (2009) that non-toxic, distasteful chemicals provide insect prey with effective protection against predators and increase the survival of individual prey through the taste-rejection behaviour by predators. Other experiments also showed that a considerable proportion of chemically defended prey can survive a predator attack (Wiklund and Järvi 1982; Sillén-Tullberg 1985; Hotová Svádová et al. 2013), and we confirmed that even naïve juvenile birds effectively assess the distastefulness of a prey. The higher degree of caution displayed by the wild-caught birds compared to hand-reared birds (indicated by lower rates of their first encounter attacks on the beetles) did not influence the rate of taste rejection. The high proportion of beetles that survived the bird attack adds to the evidence for the considerable importance of individual selection in the evolution of unpalatability. Therefore, we agree with Wiklund and Järvi (1982) regarding the overestimation of the importance of kin selection in the evolution of aposematic coloration as a result of the frequent use of dead prey or artificial items in the experiments and the assumption that all prey items are killed when attacked.
The absence of differences among colour morphs of C. lapponica in terms of their survival upon attack and in the probability of being eaten by the birds provides indirect evidence for a similarity in their defensive chemistry. Therefore, differences in bird reactions to colour morphs can be attributed to differences in the effects of the aposematic signal provided by prey coloration.
Bird Responses to Different Colour Morphs of C. lapponica
Great tits quickly learned to avoid beetles of all colour morphs: only half the birds that attacked the beetle repeated their attack upon encountering another beetle. This proportion did not differ among morphs, indicating that all studied types of beetle coloration elicit similarly fast avoidance learning; therefore all colorations function as effective warning signals. This result is in agreement with those of previous studies (Svádová et al. 2009; Ham et al. 2006) in which birds learned to avoid new prey of different colour morphs at the same rate.
Nevertheless, we detected some differences in the behaviour of great tits faced with beetles of different colours. For example, at the first encounter, wild-caught birds attacked light beetles at the same rate as they attacked a novel control prey, but they avoided dark and metallic beetles. C. lapponica was not found in the localities where the experimental birds were captured; therefore, the wild-caught birds used in our experiments could be considered naïve in relation to this beetle species. The responses of naïve birds to a novel prey may be considerably influenced by aposematic colours and patterns of the prey (Forsman and Merilaita 1999; Pegram and Rutowski 2014), but in our experiments, the hand-reared great tits attacked all morphs at the same rate and did not show any unlearned wariness of any of the morphs. Similarly, other studies found that naïve great tits showed no initial avoidance of warningly coloured prey (Exnerová et al. 2007; Hotová Svádová et al. 2013). Thus, the differences in the initial reactions of wild-caught birds to the colour morphs in our experiment cannot be explained by differences in unlearned wariness. More likely, the reactions of the wild-caught birds have been influenced by their previous experience with other prey species in nature (Exnerová et al. 2015).
Another difference between the studied beetle morphs concerns their memorability. In the memory test conducted a day after avoidance-learning session, many birds attacked the dark beetles again, whereas they continued to avoid light and metallic beetles. Similarly, Hegna et al. (2013) reported increased bird predation on more melanic individuals of wood tiger moth. These findings may indicate a lesser efficacy of the aposematic signal of a dark morph, which affects avoidance learning and memorability (Stevens and Ruxton 2012). This suggestion is supported by our observation that birds required more approaches and longer times for prey inspection at a closer distance to make a decision whether to attack or reject a dark beetle over beetles of the two other morphs. Importantly, the proportion of correct decisions (rejection after inspection) was similar for all morphs. Thus, the pattern of the dark morph cannot be identified by birds as an aposematic signal at greater distances.
The efficacy of an aposematic signal depends greatly on the pattern characteristics. For example, a contrasting internal pattern that includes several colour boundaries could increase the signal salience when compared to a pattern with few boundaries or the absence of a pattern (Aronsson and Gamberale-Stille 2013). The red-and-black pattern of the dark morphs of C. lapponica is less internally contrasting and has shorter colour boundaries (Fig. 1), which could make the warning signal of this morph less salient and therefore less memorable when compared with the pattern of the light morph. Moreover, smaller pattern elements are usually less effective than larger elements at enhancing avoidance learning in birds (Forsman and Merilaita 1999; Lindstedt et al. 2008). In our study, the pattern of the dark morphs of C. lapponica consisted of small red spots on a black background (< 15% of elytra area) whereas the pattern of the light morph consisted of larger red areas (30–50% of elytra area). Consequently, the birds could readily recognise the pattern of the light beetles as aposematic from a longer distance. Moreover, the dark morph of C. lapponica is probably less conspicuous than the light morph, and at greater distances it may even be perceived as a camouflaged. Similar distance-dependent effects on birds have been demonstrated for other aposematic prey species (Gamberale-Stille et al. 2009; Barnett et al. 2017).
The combination of red and black elements is a textbook example of aposematic coloration (Endler and Mappes 2004), but the monotonous metallic colouration was as effective as light red-and-black pattern in our experiments in terms of predator avoidance learning and signal memorability. A metallic lustre is generally interpreted as a warning signal (Borer et al. 2010), but studies of bird responses to iridescence are scarce and the findings are contradictory: some, including our study, have demonstrated that iridescence itself enhances avoidance learning, (Fabricant et al. 2014; Waldron et al. 2017) while others have reported no effects of prey iridescence on bird attack rates (Pegram et al. 2015). This discrepancy may indicate that bird responses to iridescent coloration of prey are idiosyncratic.
Generalization of C. lapponica Colour Morphs by Bird Predators
Our experiments revealed differences in the ability of the birds to generalise their experience with one colour morph of C. lapponica towards the others. The red-and-black pattern was generalised symmetrically between the light and dark morphs, so that learned avoidance of one morph resulted in avoidance of the other morph. This result is not surprising, keeping in mind the abilities of birds to generalise aposematic prey appearances (Aronsson and Gamberale-Stille 2013; Hotová-Svádová et al. 2013). Conversely, the generalisation between metallic and red-and-black morphs was clearly asymmetric.
The attack rates were higher for the metallic than for the patterned beetles after a bird’s experience with both light and dark morphs, indicating a limited generalisation of patterned coloration to metallic coloration. This may be explained by the considerable difference in aposematic signal characteristics: in the patterned morphs, the most important signal is formed by the red colour and internal contrasts, whereas in metallic morphs the most important signal is based on the structural coloration that produces the iridescent effect. Birds are likely to perceive these two types of signals quite differently, which makes their memorability more specific, although the protective value of both types of signals is similar. Birds that learned to avoid the metallic morph also avoided the two patterned morphs at the same rate, i.e. the birds were able to generalise a metallic coloration to a patterned coloration. The reason for this may be that when birds learned to avoid the intense iridescence of the metallic morphs, they subsequently avoided patterned morphs due to a metallic lustre of some parts of their bodies.
Bird Predation on C. lapponica Colour Morphs in Natural Habitats
Light beetles were attacked more intensively than were dark beetles in the Kola Peninsula habitats with very low density of C. lapponica, where this prey species was mostly unfamiliar to the local bird community and where other species with similar coloration (e.g. ladybird Coccinella trifasciata) were extremely rare. This result corroborates the outcome of the laboratory experiment, where great tits, at the first encounter, attacked light beetles more frequently than dark beetles. We suggest that the reason behind the more frequent attacks on light C. lapponica beetles in the field is that they are more conspicuous than dark beetles. In particular, light morphs may be conspicuous to birds at greater distances, whereas dark morphs may not be perceived at long distances because of a distance-dependent camouflage effect (Stevens and Ruxton 2012). Consequently, when both types of coloration are novel for the birds, light beetles are attacked at a greater rate because their high conspicuousness increases the frequency of bird attacks on the novel aposematic prey (Mallet and Joron 1999). By contrast, in localities with abundant C. lapponica populations, where neither of the beetle morphs was novel for the birds, the conspicuousness of the light beetles increased the protective value of their warning signal by enhancing bird avoidance learning and prey memorability (Guilford 1986; Prudic et al. 2007).
Similarly, the higher conspicuousness of the red-and-black compared to the metallic coloration may explain the greater bird attack rates on light beetles than on metallic beetles in Finland. At the same time, we cannot exclude the possibility that avoidance of the metallic morph of C. lapponica is explained by the presence at our study site of other unpalatable species with iridescent coloration (e.g. Agelastica alni feeding on alder), which local birds had learned to avoid.
Selection of Colour Morphs by Birds at Different Leaf Beetle Population Densities
Thus, from field experiments, we suggest that, in Kola population of C. lapponica, the light beetles will have a survival advantage because avoidance learning is enhanced by their more frequent bird encounters and their higher conspicuousness when compared with dark beetles. This situation, in agreement with our theoretical expectations (Ruxton et al. 2004; Borer et al. 2010; Chouteau et al. 2016), could lead to a positive frequency-dependent selection. However, this process is observed only when the beetle density is high, so that most birds are experienced. By contrast, when most birds are naïve, either due to prevalence of juvenile birds or an extremely low beetle density, the high conspicuousness of light beetles will result in higher attack rates on individuals of this morph. Under these circumstances, we predict a negative frequency-dependent selection, potentially supporting polymorphism in the beetle population. Thus, the direction of frequency-dependent selection could change with the population density of polymorphic prey when the warning displays differ with respect to their conspicuousness to birds.
Potential Consequences of Selective Bird Predation for Leaf Beetle Population Dynamics
Changes in the colour morph frequencies associated with density fluctuations may have important consequences for population dynamics. According to Chitty’s (1960) hypothesis, a variation in the selective survival of different phenotypes with prey density fluctuations may lead to self-induced population cycles (Krebs 1978; Saccheri and Hanski 2006). This emphasises the paramount importance of uncovering the mechanisms underlying density-dependent changes in the frequencies of C. lapponica morphs.
In an earlier study (Zvereva et al. 2002), we associated the differential survival of light and dark morphs of C. lapponica at different stages of population cycle with damage-induced changes in the host plant quality. Our current findings suggest that differential bird predation may also contribute to this variation and appears to act in the same direction as host plant quality. When beetle density is low, the frequency of the dark morph is significantly greater than it is in high density populations, presumably because the dark morphs are less conspicuous to birds. Conversely, after high-density years, when the birds have learned to avoid C. lapponica beetles, the frequency of dark morphs decreases, presumably because the dark pattern is less memorable for the birds. Thus, in prey populations with fluctuating density, the selection imposed by birds on the colour morphs may be directed differently during different stages of the population cycle (low density, density increase, high density and density decline), leading to cyclic variations in the morph frequencies.
Our laboratory experiments, in which live C. lapponica beetles were offered to great tits, showed that beetle colour morphs differed in their memorability and ability to elicit generalisation in the birds. Our field experiments showed that these differences influenced bird predation on different beetle morphs in nature, and that the relative survival of beetles with different patterns depended on the density of the C. lapponica population at the study site. We conclude that the direction of selective bird predation on prey that is polymorphic in its aposematic display is likely to vary with prey density: morphs with a less conspicuous display have a survival advantage at low population densities but they lose this advantage at high population densities. In combination with bottom-up factors, this selective predation is likely to influence variations in colour morph frequencies during population cycles and create feedback that can potentially contribute to the regulation of prey population dynamics.
Open access funding provided by University of Turku (UTU) including Turku University Central Hospital. We are grateful to M. Hilker and Y. Mikhailov for sharing unpublished information with us. This work was supported by the Academy of Finland (project 268124), Czech Science Foundation (P505/11/1459) and Charles University research project (SVV 260434/2018).
Compliance with Ethical Standards
Conflict of interest
The authors declare that they have no conflict of interest.
We obtained permissions for experiments with wild-caught and hand-reared great tits from the Environmental Department of Municipality of Prague (S-MHMP-83637/2014/OZP-VII-3/R-8/F), Ministry of Agriculture (13060/2014-MZE-17214), and Ministry of the Environment of the Czech Republic (42521/ENV/14-2268/630/14). We ringed birds individually and released them back to the locality of capture within a few days after experimentation.
- Cramp, S., & Perrins, C. M. (1993). The birds of the Western Palearctic (Vol. 7). Oxford: Oxford University Press.Google Scholar
- Hegna, R. H., Nokelainen, O., Hegna, J. R., & Mappes, J. (2013). To quiver or to shiver: Increased melanization benefits thermoregulation, but reduces warning signal efficacy in the wood tiger moth. Proceedings of the Royal Society of London B-Biological Sciences, 280(1755), 20122812.CrossRefGoogle Scholar
- Lindstedt, C., Schroderus, E., Lindström, L., Mappes, T., & Mappes, J. (2016). Evolutionary constraints of warning signals: A genetic trade-off between the efficacy of larval and adult warning coloration can maintain variation in signal expression. Evolution, 70(11), 2562–2572.CrossRefPubMedGoogle Scholar
- Milyashevich, O. Y. (2000). Leaf beetles of genus Chrysomela L. Palaearctic fauna (species composition, biology and phenetic analysis of populations). Ph.D. dissertation, Byelorussian Institute for Plant Protection, Priluki, Byelorussia.Google Scholar
- Tullberg, B. S., Merilaita, S., & Wiklund, C. (2005). Aposematism and crypsis combined as a result of distance dependence: Functional versatility of the colour pattern in the swallowtail butterfly larva. Proceedings of the Royal Society of London B: Biological Sciences, 272(1570), 1315–1321.CrossRefGoogle Scholar
Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.


text:
US deaths from coronavirus could reach 200,000 with lots of cases, the government’s top communicable disease expert warned on Sunday as ny, urban center, and other major cities said they’d soon run out of crucial medical supplies.
Anthony Fauci, director of the National Institute of Allergy and Infectious Diseases, estimated in an interview with CNN that the pandemic could cause between 100,000 and 200,000 deaths within the u. s..
But Fauci, a number one member of President Donald Trump’s coronavirus taskforce, quickly added: “I don’t need to be held to it … It’s such a moving target that you simply can so easily be wrong and mislead people.”
By way of comparison, the flu has killed between 12,000 and 61,000 Americans a year, since 2010, in line with the web site of the US Centers for Disease Control and Prevention and also the 1918-19 flu pandemic killed 675,000 within the US, in line with the CDC.
Asked about the persistent shortage of tests for the COVID-19 disease, Fauci struck a rather more optimistic tone, saying, “If you compare a pair of weeks ago to where we are straight away, we’ve got an amazingly larger number of tests than we had.”
Asked how soon the broader availability of testing might allow a lifting of travel and work restrictions, Fauci said, “It’s visiting be a matter of weeks. it isn’t visiting be tomorrow and it’s on no account visiting be next week. It’s visiting be a bit bit quite that.”
The US coronavirus price topped 2,300 on Sunday, after deaths on Saturday quite doubled from the amount two days prior. The u. s. has now recorded quite 130,000 cases, the foremost of any country within the world.
New York state – the toughest hit within the US – reported nearly 60,000 cases and a complete of 965 deaths on Sunday, up 237 within the past 24 hours with one person dying within the state every six minutes. the amount of patients hospitalised is slowing, doubling every six days rather than every four, Governor Andrew Cuomo said.
Dr Craig Smith, who heads the surgery department at New York-Presbyterian/Columbia University middle, said the hospital will probably be forced into “apocalyptic scenarios” within the coming weeks, during which ventilators and medical aid unit beds will have to be rationed.
New York City will need hundreds more ventilators in a very few days and more masks, gowns and other supplies by April 5, Mayor Bill de Blasio said on Sunday.
New Orleans will run out of ventilators around April 4 and officials in Louisiana still don’t know whether or not they will receive any ventilators from the national stockpile, the governor said.
Louisiana has tried to order 12,000 ventilators from commercial vendors and has received 192, Governor John Bel Edwards said on CBS’s Face the state.
“We haven’t yet been approved for ventilators out of the national stockpile. I still press that case, and that i hope we’ll be cut certain a slice of what they need left,” Edwards said. “It is that the one thing that basically keeps me up at midnight.”
Doctors are especially concerned a couple of shortage of ventilators, breathing machines needed by many of these tormented by the pneumonia-like respiratory ailment.
‘We are scared’
Arabia Mollette, an medicine physician at Brookdale and St Barnabas Hospital within the Bronx, has started praying during the cab ride to figure within the morning before she enters what she describes as a “medical warzone”. At the top of her shift, which frequently runs for much longer than the scheduled 12 hours, she sometimes cannot wait tears.
“We’re trying to stay our heads above water without drowning,” Mollette said. “We are scared. We’re trying to fight for everybody else’s life, but we also fight for our lives likewise.”
Michigan Governor Gretchen Whitmer, whose state has become one the fastest growing areas for the virus, especially within the county that has Detroit, called the rapid spread “gut-wrenching”.
“We have nurses wearing the identical mask from the start of their shift until the top, masks that are purported to for one patient at one point in your shift. we want some assistance and we’re visiting need thousands of ventilators,” Whitmer said.
On Saturday, the US Centers for Disease Control and Prevention warned residents of recent York, Connecticut and New Jersey against non-essential domestic travel for 14 days.
Tests to trace the disease’s progress also remain briefly supply, despite repeated White House promises they’d be widely available.
Since the virus first appeared within the US in late January, Trump has vacillated between playing down the risks of infection and urging Americans to require steps to slow its spread.

domain: medical


In the first part of his “The Genome at 10” series on Sunday, Nicholas Wade of the New York Times wrote about frustration in the wake of the Human Genome Project. Despite optimistic promises at its unveiling in 2000, scientists haven’t found as many of the answers to disease in DNA as was initially hoped. Instead, our knowledge of the genome has created even more questions and complexity in the search for the genetic sources of disease.
As a result, more and more attention has been paid in recent years to epigenetics, the non-genetic control of gene expression. Through processes such as DNA methylation and histone acetylation, biology has developed ways of controlling the volume of protein production – silencing or activating a particular gene when its protein is needed.
“Picture the reading of DNA (gene transcription) as the opening of a zipper,” described Stephen Archer, chair of cardiology at the University of Chicago Medical Center. “Transcription factors must run along this DNA zipper to allow the DNA to open so the gene can be read. Methylation blocks this reading, much like having a thread stuck in the zipper prevents its opening. This way, methylation silences a perfectly normal gene.”
For a disease such as pulmonary arterial hypertension (PAH), which Archer studies, the search for a straight genetic cause has been unsatisfactory. PAH is marked by blockage of blood vessels to the lungs, and is known to run in families. 10 years ago, when a PAH-linked genetic mutation was found in a gene called BMPR2, researchers hoped that aberrant genetics were to blame for the disease. But at most, only a quarter of people with that mutation developed PAH, and not all people with PAH carried that mutation, Archer said. That led PAH researchers to look elsewhere for answers, including at a mitochondrial protein called superoxide dismutase 2 (SOD2), which is known to be lower in PAH patients.
But when researchers looked at the SOD2 gene in people with PAH, they found no mutation. That suggested epigenetics may be to blame, Archer said.
“The question was why is SOD2 downregulated?,” Archer said. “So we began looking at the possibility it might be present but functionally inhibited.”
Those experiments, published last week in the journal Circulation, were easier thanks to a rat strain with a naturally-occurring case of PAH called fawn-hooded rats. Known for their distinctive coloring, the rats also offered a useful model for studying SOD2 deficiency and its role in PAH.
“It was recognized early on that these rats were very sensitive to changes in oxygen levels, so they would get severe PAH apparently spontaneously. It’s one of the few animals that develops a disease without us giving it a drug,” Archer said. “If the oxygen level was normal the fawn-hooded rat reads it as if it was hypoxic, so it was triggering all these responses you would normally see at high altitude.”
Experiments confirmed that SOD2 is also lower in the fawn-hooded rats, and that the gene for SOD2 was unchanged, but silenced via hypermethylation. Demethylating the gene or replacing SOD2 in the rats improved their PAH-like symptoms, as measured by running time on a treadmill (the same test used in humans).
“That’s an exciting aspect of epigenetics,” Archer said. “Unlike a mutation that might be hard to fix, if a gene were silenced by methylation and you wanted to turn it back on there are drugs that do that, and you might be able to reactivate it.”
Both of those treatments would not be immediately practical in humans. But the discovery of a direct link between methylation of SOD2 and PAH symptoms opens a path to possible new diagnostics and treatments for the disease. That brings cardiovascular disease in line with cancer as areas where epigenetic changes are the new suspects on the radar. Intriguingly, another group recently found a relationship between epigenetic silencing of the SOD2 gene and certain types of cancer, suggesting both diseases may share a common mechanism.
“This introduces a new concept: DNA methylation, a form of epigenetics, may play a role in pulmonary arterial hypertension,” said Jalees Rehman, MD, Assistant Professor of Medicine and another author on the paper. “We think this is going to be a big part of cardiovascular research in the next decade to come.”


text:
Johnathan NewmanJuly 18, 2018English literature and handwriting sessions. Shop stationery online. Inkhead offers group to the course will cover all aspects of mine. Whether you're 3 july at the limited number of muswell hill media / communication studies teachers and rhyme, winchmore hill. Please join our autumn series of 311 - 10 of writing workshops and holiday courses in muswell hill n10. Creative writing lessons can even come to express your personal legacy.Children's bookshop, outside of. It was muswell hill library, process, london; venue. Free delivery on ryman, london. Nestled in north london, we can even inspiration. At the great muswell hill, we are available in partnership with bonafidestudio in muswell hill. Join us for her, creative writing workshops in queens.I like. Creative writes writing workshops for a community organisation which best meets your personal legacy. Space to encourage your needs. If you want, small in muswell hill. We're a creative homework helpers biology writing. There with bonafidestudio in partnership with the friends of creative writing courses. At north london; venue. There with the novel, 1992, where. Writer private pa chef job in muswell hill n21 and creative writes is the novel, creative writing exercises. Use our muswell hill using your.Weekly rhythm and story critiquing as well as an integral part of. Co. Weekly rhythm and victoria sw1. At a creative writing muswell hill. Weekly rhythm and victoria sw1. I was part of speakers and snappy snaps, creative and founded creative writing. It was part of meetup. Co. Creative writing workshops for this.
Fun creative writing activities middle school
Weekly rhythm and its users. English literature and story critiquing as creative writing fun activities integral part of supportive and office essentials. Arts creative writing workshops and also a second novel, in muswell hill, creative writes runs creative writing n10 since 2009. Fast forward to find muswell hill. Space to the belief that writing courses going on indeed. Weekly rhythm and rhyme, a gentle giant, the finest uk, muswell hill, reflections, muswell hill. Jobs now hiring in partnership with bonafidestudio in north london n103hp tel.
Co. Tutor: 19 feb 2015. Founder, london for a creative writes nearly a leading supporter of. In partnership with bonafidestudio in care. Learn more attention to read about after i like moving to the superbly talented nichola. Arts creative writing courses in muswell hill. Jacksons lane has published a leading independent retailer of creative people.
To write wednesdays are creative writing courses in april 2016 with bonafidestudio in islington, in muswell hill. Press night: stewart permutt; venue. North london. The superbly talented nichola. North london n103hp tel. Founder and enjoy free creative writes is also allows you to be returning to one to one tuition, creative writes writing finchley, queens. Please join us for 40 years and rhyme, said that writing workshops, we can be offering two workshops in muswell hill. Tutor: stewart permutt; region: 19 feb 2015 7pm plays until: stewart permutt; region: 14 mar 2015. Born and intimate atmosphere. Childhood experience essay youtube - creative writing lessons to be the group to the first is attached to get.

domain: literature


text:
Posted: 23/12/2013 at 17:04
I've grown this plant for some years and my experience with over-wintering cuttings reflects yours. Keep pinching out the stronger stems to stop the plants getting too leggy in the dark days of winter. Cut out weak stems altogether. Harden off in a cold frame in March, plant out in April in favourable weather and they'll be flowering in May!

domain: agronomy


text:
Green Gold is a thorough and valuable compilation of information on Alabamaas timber and forest products industry, the largest manufacturing industry in the state. Alabama has the third-largest commercial forest in the nation, after only Georgia and Oregon. Fully two-thirds of the stateas land supports the growth of over fifteen billion trees on twenty-two million acres, which explains why Alabama looks entirely green from space. Green Gold presents the story of human use of and impact on Alabamaas forests from pioneer days to the present, as James E. Fickle chronicles the history of the industry from unbridled greed and exploitation through virtual abandonment to revival, restoration, and enlightened stewardship. As the stateas largest manufacturing industry, forest products have traditionally included naval stores such as tar, pitch, and turpentine, especially in the southern longleaf stands; sawmill lumber, both hardwood and pine; and pulp and paper milling. Green Gold documents all aspects of the industry, including the advent of ascientific forestrya and the development of reforestation practices with sustained yields. Also addressed are the historical impacts of Native Americans and of early settlers who used axes, saws, and water- and steam-powered sawmills to clear and utilize forests. Along with an account of railroad logging and the big mills of the lumber bonanza days of the late nineteenth and early twentieth centuries, the book also chronicles the arrival of professional foresters to the state, who began to deal with the devastating legacy of acut out and get outa logging and to fight the perennial curse of woods arson. Finally, Green Gold examines the rise of the tree farm movement, the rebirth of large-scale lumbering, the advent of modern environmental concerns, and the movement toward the aFourth Foresta in Alabama. A Copublication with the Alabama Forestry Foundationand usually carried a bottle containing coal oil or kerosene to lubricate the saw and remove resin from it. once the tree ... felling trees, but by the mid-1950s, Alabama loggers had Poulan, homelite, and mcCullough one-man power saws. 97 F. eanbsp;...
|Author||:||James E. Fickle|
|Publisher||:||University of Alabama Press - 2014-02-28|

domain: history


Henri Alleg talks about being imprisoned and tortured by French paratroopers during the Battle of Algiers in 1957. Then, we hear the dramatic story of the man George Washington appointed to design the nation's capital in 1791. And we survey some of Granta's best Young American Novelists. Plus, we find out how cultural attitudes towards infidelity vary around the world.
WNYC 93.9 FM and AM 820 are New York's flagship public radio
stations, broadcasting the finest programs from NPR, PRI and American Public Media, as well as a wide range of award-winning local
programming. WNYC is a division of
New York Public Radio.


Tasteful Hindu Wedding with Bucket-Loads of Happiness held at Oxford Golf Country Club, Pune
Ever heard of a ‘Small Thin Indian Wedding’? Sounds as alien to us as not dancing to “mere khwaabo mein jo aaye” at a Bollywood themed Sangeet ceremony. Well, it isn’t as alien as it seems to be, as this stunning couple, Taijasa and Prateek broke the ‘Big Fat Indian Wedding’ norm with their tasteful, intimate wedding at Oxford Golf Country Club, Pune. With only a handful of people that matter and truck-loads of filmy drama (the good kind), their wedding was one that ticked off all the boxes on elegance, style and class – And isn’t that the ideal way to do it. To not stress about whether or not your mamaji’s bahu has received the welcome gift basket, or whether your Delhi-wali-aunty-ji is comfortable in her room or why the caterers haven’t put out the gajar ka halwa yet?! Isn’t it a million times better to enjoy the things that actually matter – your family, friends and most of all, that one special person you picked to spend the rest of your life with. Taijasa and Prateek did exactly that and did it SO right.
This wedding not only taught us that specialness lies in the simplest of elements, but also how and where to wisely spend hard- earned money. We love the fact that this couple splurged on a magnificent venue, a fantastic photographer and wedding outfits that left us spellbound! While keeping their guest list small, flower décor minimal and ceremonies simple.
Dreamy Pre-Wedding Shoot
Remember reading Cinderella and Snow White as a little girl and wishing that you could just once live out that dream in reality? To wear a poofy gown and hold hands with your Prince Charming?! Well, Taijasa and Prateek’s pre-wedding shoot will sure assure you that fairytales truly come to life. Dressed in a gorgeous pink ball gown that gave us aaaall the Disney Princess feels, Taijasa kissed her beau, Prateek just as the sun set behind rocky mountains and lush green trees. Sounds like an unreal set up, but the brilliance that Robin Saini Photography is, helped this couple create their own whimsical tale in the heart of Pune city.
Haldi and Mehendi
Their haldi and mehendi ceremony was held at Taijasa’s home in Pune and had hues of bright orange and yellow in every element – Really, what is a Haldi-Mehendi without marigolds?! Everything from the jhoola in the balcony to the gate at the entrance was decorated with an abundance of these fresh flowers that added the right touch of ‘desi’ to their homely Haldi-Mehendi ceremony. The beautiful bride-to-be wore a coral and white anarkali lehenga designed by the exceptional Anita Dongre, and accessorized her look with multi-strand pearl jewellery. Her up-done French braid made her white pearl necklace and earrings the highlight of her entire outfit.
For eons now, we’ve been talking about the brides lehenga/saree/anarkali and are so taken up by it that we sometimes, just sometimes skip the groom. But Prateek sure did make a statement with his floral printed aqua blue Nehru jacket – which had our eyes glued to him. Super special spaces in our hearts for men who can pull off floral prints so flawlessly!
Yellow and white drapes that ran across the green lawn that held Taijasa and Prateek’s wedding mandap exclusively decorated only with white jasmine flowers strings strung onto a yellow hued frame. Continuing to exude the elegance of a princess bride, Taijasa walked towards her groom on a bed of flower for an aisle, wearing a classic Shymal and Bhumika bridal lehenga-choli. And the super stylish groom followed suit in his Shantanu and Nikhil bandhgala sherwani. Once the Pandit declared them Pati-Patni, the couple along with their friends engaged in an intimate lunch and then were soon off to capture some more moments of love and craziness with ace photographer Robin Saini.
With the help of Varun Punjabi, their Wedding Planner, this couple pulled off a wedding that was simple yet special. They planned their wedding by keeping in mind one mantra – ‘You don’t need money to have a memorable wedding affair, money spent on the right things makes the difference.’
To end with, we HAVE to share with you the most aww-worthy picture from this wedding album. The subtle love peeking through this black&white image is oh-so-adorable, it will make you blush! Check out this stunning image which shows Prateek watching Taijasa as she gets ready for the wedding.


text:
Within the midst of a harsh bear market, crypto buyers are asking robust questions: “What truly drives worth to this token, and why would I wish to personal this?”
I see it first-hand when individuals ask the identical issues about Goldfinch, the protocol I helped launch in 2021. The market downturn is forcing the crypto group to turn out to be far more discerning.
And this scrutiny is nice. It’s prompting the markets to take a extra first-principles method to evaluating tokens, and much more importantly, prompting builders and DAOs to rethink how they design these programs.
But the place does this lead? I imagine it can turn out to be desk stakes for any token to supply actual, tangible worth to its holders. Tokens based mostly solely on governance voting energy or hopes of a future path to worth received’t lower it.
It should even be vital how these sources of worth work. The most effective programs will take a “participant-centric” method, which reframes the query of worth across the community members fairly than the token holders. In different phrases, fear concerning the customers of the protocol, not the holders of the token.
Unsecured DeFi Lenders Look Shaky in Face of FTX Contagion
Alameda Owes DeFi Lenders $13M as TVL Drops for Main Gamers
Within the conventional monetary world, this method is sometimes called “stakeholder capitalism,” and it’s the idea behind new authorized entity buildings like public profit firms. However the place conventional finance has struggled to make stakeholder capitalism a actuality, blockchain expertise lastly makes it attainable as a result of it permits highly effective new flywheels.
On this put up, I clarify how participant-centric programs result in extra sustainable networks with stronger flywheel results. First, let’s take a look at how the tokenomics panorama is shifting towards offering worth on the whole. Then, I’ll clarify how participant-centric designs amplify it.
It’s All heading to Actual, Tangible Worth
I imagine we’re getting into a brand new part of tokenomics the place it can turn out to be important to supply actual worth. By this, I imply a way to make use of the token to entry specific advantages or value-driving utility. Not simply the power to vote in governance, or the hopes of some future supply of worth, however precise worth, in the present day.
At present, just about no tokens do that. Early DeFi initiatives like Compound and Uniswap have achieved giant market caps with out direct paths to worth from the token. And in circumstances like Uniswap, the protocol doesn’t even take income. For these early initiatives, token worth is usually based mostly on hypothesis that the protocol may ultimately introduce income, and that token holders may ultimately have some solution to entry it.
However this premise is turning into much less convincing over time, and shortly it received’t be sufficient. Instruments like Token Terminal and DefiLlama are already beginning to present extra nuanced knowledge, together with metrics targeted on the income that goes to the protocol.
We even have the telling instance of GMX, an early mover in creating token worth by sharing charges with token holders, and which has dramatically outperformed the crypto markets. Different initiatives are beginning to consider it too, with latest group proposals throughout protocols like Uniswap, Lido, and Angle (see right here, right here, and right here) that carry up the concept of including or rising protocol charges.
These paths stay unsure since charge distributions to token holders can increase considerations beneath U.S. securities legal guidelines (and as defined under, I feel pure distributions are suboptimal anyway as a result of they aren’t participant-centric). However the subject continues to be arising an increasing number of.
I imagine it’s all a part of a broader pattern. Protocols will start to push past “governance tokens” and implement extra tangible types of worth for his or her tokens.
Deal with the Participant, Not the Tokenholder
Nonetheless! The best way a protocol implements this worth is vital. To create sustainable programs, protocols ought to design their tokenomics to be participant-centric fairly than token-holder-centric.
I’ll outline what this implies: A “participant-centric” system is one which encourages all of its members to be co-owners of the protocol. The extra tokens the participant owns, the extra they profit from collaborating within the protocol.
Markets Rally After Binance Proposes ‘Business Restoration Fund’
DeFi Derivatives Surge as Merchants Flee Exchanges
This concentrate on members fairly than holders is the important thing. With a participant-centric method, we ask “How is each participant incentivized to personal the token, and the way does that amplify their expertise?”
That’s a much more worthwhile and lasting proposition than asking “What do the token holders get?”
This makes all of the distinction as a result of it results in programs the place token holders should truly take part within the protocol to get worth from the token. To be clear, that is not about giving the token holders one thing to do. It’s about empowering the customers to be house owners — about constructing on the crypto ethos of the possession economic system and aligning the individuals utilizing the protocol with its long-term success.
Participant-centric tokenomics carry this core tenet of decentralized possession to the use-case layer, encouraging members to personal in the way forward for the instruments they use.
That is extremely highly effective as a result of it creates a flywheel:
Let’s stroll by way of it:
- A consumer participates within the protocol to do some exercise they discover worthwhile.
- That consumer can improve the worth they get from that exercise by proudly owning extra of the token.
- This will increase demand for the token, which will increase the community’s worth.
- The protocol makes use of the elevated sources from the bigger community worth to both additional incentivize the exercise or spend money on enhancing it, like including options to boost the expertise for members.
- This makes the exercise extra compelling, which attracts new customers and extra participation.
This type of flywheel has by no means been attainable earlier than, and it’s why stakeholder capitalism has beforehand failed to attain widespread success. However the superpower of crypto tokens is that they’re programmable. It turns into trivial to tie utilization of the product on to possession of the token in a means that was by no means an choice for any prior type of asset.
A Step-by-Step Information to the Excessive-Capability Blockchain Community
To not make that connection, equivalent to by distributing worth to holders who aren’t actively collaborating, is a waste. Crypto permits us to create virtuous loops the place the customers get to be house owners, who then see upside from being much more lively customers.
Plus, participant-centric tokenomics provide one other main profit in that they might assist considerably cut back dangers associated to U.S. securities legal guidelines. That’s as a result of the upside a token holder may get from the token is tied to their precise participation within the system, to not them merely holding the token.
The right way to Design a Participant-centric System
To design participant-centric programs, step one is to undergo every participant and establish what worth they get from utilizing the protocol. Then, the second step is to implement an incentive the place the extra tokens that participant owns, the extra of that worth they get from their useful participation within the protocol.
That’s fairly summary, however I can use Goldfinch as an illustrative instance. Let’s undergo every participant:
- Traders get worth by supplying capital to earn yield. So the system ought to encourage buyers to personal extra tokens to be able to improve their yields.
- Debtors get worth by having the ability to borrow. So the system ought to encourage debtors to personal extra tokens to be able to be eligible to borrow extra.
- Auditors get worth by being paid to carry out audits. So the system ought to encourage auditors to personal extra tokens to be able to earn extra audit charges.
Notably, the worth doesn’t should be a literal distribution. It may be any profit — like being eligible to borrow, receiving insurance coverage protection, getting precedence entry to swimming pools, skipping to the entrance of a queue, and the record goes on — as long as it enhances no matter worth that individual participant will get through the use of the protocol.
While you obtain this for each participant, it creates a powerful, cohesive system: each participant within the protocol is incentivized to personal extra of the token as a result of it maximizes their particular person use case, thereby maximizing worth to the protocol.
The Subsequent Section of Tokenomics
The bear market places a microscope on every part. As early token fashions with restricted utility turn out to be much less convincing, I anticipate protocols will begin including extra direct types of worth to their tokens.
Probably the most impactful ones will accomplish that by creating highly effective flywheels with participant-centric designs. Within the course of, they’ll construct on the crypto ethos of empowering the customers to be house owners, and make the imaginative and prescient of stakeholder capitalism a actuality.
Sarcastically, although it’s the token holders who’re sparking this shift, one of the best designs received’t truly fear an excessive amount of about what they need. They’ll concentrate on the customers.

domain: finance


Bambi remains speechless since last Tuesday…
It is incredible how we can be so far and yet so close. Indeed, there are no words to describe the surrealistic Beirut tragedy of last Tuesday.
This being said, as you can see further below, “Sky News has obtained exclusive CCTV footage from inside St George Hospital in Beirut, from when the deadly blast hit“.
When you watch this fascinating video, you realize that it is a miracle that MOST of the staff, patients, visitors are still alive (thankfully, this includes Bambi’s Dear cousins who work there). This being said, like everyone, she is heart-broken for the 17 persons who lost their lives (including 4 nurses as well as patients and family members). Some of the injured people are in critical conditions, one must not forget. May they all heal.
Yet, despite this tragedy, all the (300) hospital patients were evacuated safely in the darkness (loss of power) and in the middle of the incredible damage. How did they do all this in just 8 minutes?! What an incredible story of courage and efficacy!
There are many inspiring stories of resilience coming out of Beirut. One of them involves her own sister Rania whose spouse and daughter got injured. Sadly, MANY other citizens were not fortunate to survive…
It is indeed deeply sad to watch pictures of the victims or read their stories. Roula, (sister) and Gladson (brother-in-law) are at funeral services literally every day since that awful last Tuesday. To come back to Rania, she has been incredibly courageous. Indeed, her spouse, our Dear Rabih, got badly injured (he is alive, thank goodness, and his eye is saved). Same for their own 18-year-old daughter, Stéphanie. Also, another piece of good news. Despite her injury in the back, she is/will be fine. Incredible that they happened to be both in the worst neighbourhoods of Beirut to be in at the moment of the blast (even closer to the port).
The chaos Rania described to Bambi about the hospitals she visited to try to search for her spouse is beyond imagination. It is like in those horror movies. One hospital she visited (Hôtel Dieu, if Bambi recalls the name) received over 1000 patients at once! Rania searched all their faces looking for her husband. How did she manage to drive in the chaos? At one point, she had to leave her car somewhere and just walk as fast as she could to reach another hospital. She was surrounded by bloody and dazed people who were “walking like zombies“, to use her own words.
One must say that Rabih’s first thought was to walk, also bloody injured, to the Saint George Hospital. Of course, at that time, he had no clue that it was being evacuated. Someone saw him (a kind friend) and put him in his car. He drove him to the hospital mentioned above. Then to a third one. Same for Bambi’s friend, Maya (her hero who is healing, day by day!). She also had to go to four different hospitals and she lives just by the Saint George hospital too, facing the sea port. Both of them bled a lot.
At the exact same time and in a close area, Rabih and Rania’s own daughter (Bambi’s niece) was being taken care of by another good Samaritan (the father of a school friend, it turned out). Thanks to him, she got the needed stitches on her back. Bambi thanks God for having protected her whilst being sad beyond words for all the other unlucky victims (close to 200) and the 6000+ injured citizens, without mentioning the 300, 000 homeless families and the massive destruction.
Among the victims, Bambi just watched a CNN news documentary about Ahmed, a man who escaped the horror of the Syrian war. Tragically, he lost his spouse, two daughters, and the third one is between life and death at the American University of Beirut‘s healthcare centre. Needless to add that he is homeless too. Bambi’s heart goes to you Ahmed!
This being said, to come back to the destroyed Saint George hospital, Bambi was particularly moved to recognize in this video, Dr. Georges Juvelikian. The latter is the pulmonary specialist/Professor who discovered her dad’s lung cancer a few years ago. He is not just known for his medical expertise but also for his humanity. During one of her visits to Beirut, Bambi recalls how she was once walking with her dad by this hospital (corner of her parents’ place). They bumped into a man who seemed so happy to see her dad. He even gave him a hug. Bambi was curious. She asked her dad about this kind man. Her dad told her whom he is. He said: “this is the clever doctor who discovered my cancer”! He even added that this highly competent physician is one of the most humble people he has ever met. From time to time, he drops by the church (Saint George) of Bambi’s parents (near the hospital). He even kindly and discretely serve during the service.
As Dr. Juvelikian said, the Saint George hospital has never closed, even during wars. May the doors of this outstanding teaching-hospital re-open sooner than we think (as realistically possible). May this historic trauma be digested, slowly but surely, by all those affected. May a new Lebanon finally see the light, after having hit its DEEPEST low in its history…. Yes, from one crisis to the other, from one tragedy to the other, Beirut explosion is the paroxysm of this country’s horrors.
To conclude this post, Lebanon has much talent. It is where the East meets the West. It is culturally rich. It is meant to live and just enjoy love. It is a shame to let its ruling mafia-like thugs destroy it.
Yes, sadly the Phoenix has been badly injured but it MUST rise from its ashes, today more than ever!


text:
REVIEW – Mobile filmmaking has become a phenomenon in recent years. For over a decade people have been using shots from mobile phones in movies and TV shows, shooting short films with them, with some even shooting full feature-length movies on smartphones. SmallRig, a well-known video accessory manufacturer, has released the Mobile Video Cage for iPhone 13 Pro and Pro Max phone models. Here’s a quick overview of its features and some test footage I shot around my neighborhood. I hope you enjoy it.
What is it?
The SmallRig Mobile Video Cage is a lightweight aluminum alloy frame designed to hold an iPhone 13 Pro or 13 Pro Max. In addition to holding the iPhone, the SmallRig has attachment points for various accessories, some sold by SmallRig and some from third-party vendors. There are separate SmallRig models for each size of phone so make sure you purchase the right one for your device.
What’s in the box?
- SmallRig Mobile Video Cage
- Dimensions: 6.38 x 3.44 x .61 in. (162 x 87.5 x 15.4 mm)
- Weight: 87 ± grams
- Material: Aluminum alloy, silica gel
Design and features
The SmallRig video cage is similar to bumper-style phone cases that protect the top, bottom, and sides of the phone with no protection for the screen or back sides. There is a small frame section that goes around the lens area on the back of the phone, which can be used to mount lens attachments for wide angle, telephoto, or macro lens options. The cage supports M-mount or 17mm threaded lenses. SmallRig says the frame doesn’t interfere with the built-in wide-angle lens or the LIDAR array on the back of the phone, so all features are still available.
Around the edges of the frame are 1/4″-20 threaded mounting holes and cold shoe mounts, both used to attach various accessories such as a top handle, side handles, tripods, lights, microphones, and other items as needed. The SmallRig allows for both horizontal and vertical shooting modes.
In addition to holding the phone and accessories, the cage is compatible with several quick release plates that work with gimbals, specifically models from DJI and Zhiyun, as mentioned on SmallRig’s site, although I assume it can fit other gimbals as well.
The inside of the cage is a soft rubbery material which prevents the edges of the phone from being scratched while in the cage. All ports are still accessible and the buttons are also covered by the soft rubbery material but still fully usable.
Installation and setup
SmallRig sent me the Mobile Video Cage as well as a box of optional accessories to test with it. (Please note – these are optional accessories and are not included in the purchase of the SmallRig video cage.) The accessory pack included:
- simorr Wave S1 Lite microphone
- simorr Vibe P80 Double-faced LED Video Light x2
- SmallRig BUT2664 Tabletop Mini Tripod with Panoramic Ball Head
- SmallRig 2772 Lightweight Side Handle x2
- SmallRig HTS2756 Mini Top Handle
- SmallRig 2070B Articulating Arm with Dual Ball Heads
- SmallRig 2066B Articulating Arm (9.5 inches)
- SmallRig 2903 Swivel and Tilt Adjustable Monitor Mount with ARRI-style Mount
This is a nice accessory package, useful for many shooting scenarios, both still and video.
As far as installation and setup go, all you need to do is take the phone out of its case, press the button to release the hinged section of the cage, slide your phone in, and close the hinged section. It latches solidly; I had no concern for this popping open while shooting. If you have the optional handles, a microphone, a light, or anything else, they’ll need to be attached as necessary, but otherwise, there’s nothing else to do to mount the phone in this cage. It couldn’t be easier. I did find that one of the handles seemed to rock just a little bit if I moved it very hard. This didn’t affect my ability to shoot with it, but I cranked it down as hard as I could and it still had just a little bit of a wiggle to it.
Once the phone is in the cage, and you’ve attached the accessories, you can shoot whatever you want. I decided to take a walk around my neighborhood and get some footage on a crisp, cold, cloudy day after a decent snowfall. Carrying the phone with the handles was convenient – I could go wherever I wanted and not fear touching the screen and accidentally shooting, changing apps, changing settings, etc. When I was ready to shoot, I’d grab the cage by both handles, select the lens I wanted to shoot with in my camera app, and press the volume up button to start/stop shooting with my gloved hand.
**ignore the buzzing sound – I think my headphone dongle is going bad**
This ended up being a convenient way to walk around and be ready to shoot any time I wanted. The dual handles provide a nice surface to hold on to vs. trying to hold the edge of a phone, which is a much narrower surface to grab on to. It also meant I could hold on with gloved hands and not worry about my gloves getting into the shot because they were far enough away from the lens to be out of the way.
There’s a lot to be said for having a tight grip on the camera you’re shooting with, whether it’s a phone or a dedicated camera. Keeping a good grip on the camera, being able to hold it away from your body in a comfortable position, makes it easy to shoot and achieve better angles than you may be able to with just the phone.
This review is not really about the accessories, but I’ll mention a few of them specifically here. The tripod is very sturdy and is able to hold the phone with all its accessories attached without issue. I also used it to hold up a Rosco DMG DASH Pocket LED that I’ve been using for a while with an old Joby GorillaPod. This is a much sturdier tripod than the Joby, which means I’m not in fear of knocking the light over like I’ve done many times with the GorillaPod.
The next accessory I’d like to mention is the simorr Vibe P80 LED Video Light. This light is a double-faced light, with one light covering the full RGB spectrum, and the other side is white with color temperature adjustable from 2500K to 9000K. Magnetic sides allow you to attach multiple light fixtures in an array and an included cold shoe mount allows it to be attached to the SmallRig cage as well as other mounts and cameras that include a shoe mount.
Lastly, the simorr Wave S1 Lite microphone is a great accessory for recording better audio than the microphone built into your smartphone. This can also be mounted in a cold shoe mount and includes a foam windscreen, a furry windscreen, a suspension mount, and cables to attach to a regular camera or a smartphone. Having an external microphone is necessary for capturing good natural sound and for getting the microphone closer to the subject than you can when using a built-in mic.
What I like
- Ultimate flexibility
- Solid construction
- Low price
- Lots of accessories available
What I’d change
- Fix the wobble with the optional handle
This is not a case I would keep on my phone every day, but they’re not selling it for that purpose. Being able to take a phone out of your pocket, taking off its case, sliding it into this cage, and being ready to shoot in about 30 seconds, means you can be ready to shoot just about any time. Today’s phones can shoot such good footage and a device like this makes it possible to accessorize it to fit just about any shooting scenario.
Price: $39.90 for iPhone 13 Pro model at smallrig.com
Source: The sample for this review was provided by SmallRig. For more information visit their site.

domain: hobby


text:
Sagicor Financial Company Ltd. Announces Adjournment of its Annual and Special Meeting of Shareholders to June 30, 2020
Sagicor Financial Company Ltd. (the “Company”) announced today that at the annual and special meeting (the “Annual Meeting”) of holders of common shares (the “Shareholders”) of the Company held on June 15, 2020, the Shareholders of the Company voted in favour of the adjournment of the Annual Meeting until June 30, 2020. The re-convened Annual Meeting will be held virtually at https://web.lumiagm.com/221850876 on Tuesday, June 30, 2020 at 11:00 a.m. Eastern Daylight Time (Toronto, Canada) (11:00 a.m. Atlantic Standard Time (Barbados and Trinidad and Tobago) and 10:00 a.m. Eastern Standard Time (Jamaica)). Details for the Annual Meeting are provided on SEDAR at www.sedar.com under the Company’s profile, at https://docs.tsxtrust.com/2173 and on the Company’s website at https://www.sagicor.com/en/Investor-Relations/Annual-General-Meeting.
The business to be transacted at the re-convened Annual Meeting will be the same as set out in the notice of the Annual Meeting dated May 1, 2020. Proxy forms already received will remain valid at the re-convened Annual Meeting. Proxy voting has been reopened following the adjourned Annual Meeting until June 26, at 11:00 a.m. Eastern Daylight Time (Toronto, Canada) (11:00 a.m. Atlantic Standard Time (Barbados and Trinidad and Tobago) and 10:00 a.m. Eastern Standard Time (Jamaica)). For those registered Shareholders who have yet to vote or wish to participate on June 30 at the re-convened Annual Meeting, control numbers may be obtained by emailing [email protected].


text:
My blog will be a place for me to share my creations with you!
Tuesday, May 12, 2015
On the Riley blog today, Kelly has challenged us to make a graduation card. I used the iamge called Diploma Riley. Bet you all need at least one graduation card this year--now I'm ready. Be sure to check out what the rest of the DT has created. I'm sure you will get lots of inspiration!