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Ecological correlates of sociality in Pemphigus aphids, with a partial phylogeny of the genus
Nathan Pike1,
John A Whitfield2 &
William A Foster2
Because the systems of social organisation in the various species of Pemphigus aphids span the continuum from asociality through to advanced sociality (typified by the possession of morphologically specialised soldiers), the genus is an ideal model clade in which to study the influence of ecology on the origins of eusociality. We made detailed study of the ecology of three gall-dwelling species that show clear differences in their levels of social behaviour. To elucidate evolutionary relationships and to attempt to estimate the number of origins of sociality, we also created a phylogeny based on sequences spanning the mitochondrial genes Cytochrome Oxidase I and II for nine species of Pemphigus.
P. spyrothecae, a highly social species with aggressive morphologically-specialised soldiers, has the longest galling phase, unsynchronised development of a large number of individuals in a densely-populated gall, and an extended period over which alates emerge. P. populi, a species with no soldiers, has the shortest galling phase, synchronised development of a small number of individuals in a sparsely-populated gall, and an extremely brief emergence period. The ecology of P. bursarius, which has behavioural soldiers that are not morphologically specialised, is intermediate between these two extremes. The galls of P. spyrothecae and P. bursarius form small openings during the course of the season and predation-related mortality is relatively high in these two species. Conversely, predation does not occur during the galling phase of P. populi, which has no soldiers but makes an entirely-sealed gall.
The phylogeny allowed us to infer one likely point of origin of basic social defence and two independent origins of enhanced defence. Based on current knowledge of behaviour, the phylogeny also suggests that the defence trait may have been lost at least once.
The life-history strategy of P. spyrothecae appears to be geared towards defending the colony against the constant threat of predation that faces the inhabitants of a long-lived, open gall. The life-history strategy of P. populi, on the other hand, is to avoid predation in the closed gall fortress and flee for the secondary host at the earliest opportunity. The life-history strategy of P. bursarius appears to represent a compromise between these strategies.
The transition from solitary to social living is one of the key events in evolution: the rarity of this transition has provided a classic problem for evolutionary biologists and the unparalleled success of social animals is of central interest to all ecologists. To investigate the factors underlying this transition, it must be studied in a phylogenetic context and we therefore require clades of extant species that have several independent origins and different degrees of sociality. Such clades are exceptionally rare. For many groups, the number of independent origins of sociality is very small, these origins may be buried deep in the past, and the phylogeny may not be known. Other groups, for example bees and thrips, are more promising. The social aphids provide particularly fruitful material because there have been many independent origins of sociality and there are clades of known phylogeny containing species covering the spectrum from solitary to fully social organisation. There are estimated to be at least 17 independent origins of sociality in aphids (T. Fukatsu, pers. comm. 2006) even though only 1% of aphid species is social [1, 2]. This dwarfs the equivalent estimates for other social taxa (e.g. 1 origin in the ants [3], 1–4 origins in the termites [4], fewer than 11 times in the wasps and bees [5]). In addition, aphids have a genetic system – clonality – that is virtually unique among social insects. This reduces the complexity caused by genetic conflict that is present in most social systems and allows us to concentrate on the role of ecological factors, which have probably driven the evolution of all animal societies [6].
The vast majority of aphid species simultaneously produce more than one morph, and the different morphs routinely have different direct reproductive fitnesses. For example, wingless morphs may produce more offspring than winged morphs [7]. It is certainly true that such reproductive sacrifice may constitute an altruistic act which enables the altruist to contribute indirectly to the fitness of its clonemates though behaviours such as migration. However, in line with the approach most commonly taken, we exclude such altruism from our definition of true aphid sociality, which we consider to have occurred only when a species has acquired soldiers, morphs that actively defend against predators. It is undoubtedly relevant that these soldiers are often the agents of socially advanced behaviours other than defence, such as nest cleaning [8, 9], nest repair [10, 11] and invasion and exploitation of alien colonies [12].
Every species of social aphid forms and inhabits a plant gall at some point in its life cycle [13]. The galls provide a rich and indispensable resource to the aphids which are therefore classic examples of "fortress defenders" [14]. The gall is especially important for serving as a physical barrier to predators and to alien aphid invaders that seek to exploit the altruism of the home clone, disrupting the alignment of genetic interests among colony mates.
This study concerns the aphid genus Pemphigus which contains species that exhibit almost the entire spectrum of sociality from the total absence of defenders to the presence of aggressive, morphologically specialised soldiers [15]. Pemphigus soldiers generally occur in the first instar of the virginoparous morph on the primary host but defensive behaviour also continues to a lesser degree in later virginoparous instars [15]. It is clear that the gall is particularly crucial to the social biology of Pemphigus aphids because soldier morphs have been found only at the galling stage [16]. We produce a partial phylogeny that includes nine of the sixty-five species of Pemphigus that have been described [2]. The species of the genus are spread across the Northern Hemisphere and many have cosmopolitan distributions [17]. Populus (poplar) is the primary host on which the galls are formed [18] and host alternation is the norm, but not the rule: three of the 17 species that have been sufficiently studied spend their entire life cycle on poplar. Because the genus is entirely without secondary-host soldiers, we can exclude the possibility that a second, dissimilar environment influences social evolution and we can thus focus solely on the galling phase. The Pemphigus species that have been scrutinised for the presence of soldiers are presented in Table 1. In this study, we have examined in detail the natural history and ecology of three species: Pemphigus populi, which does not have soldiers; Pemphigus bursarius, which invests moderately in gall defence using monomorphic nymphs that behave as soldiers; and Pemphigus spyrothecae, which invests extensively in morphologically specialised soldiers.
Table 1 The species of Pemphigus in which soldiers have been sought
Within the genus Pemphigus there is known to be considerable variation in factors such as time spent on the primary host, the number of aphids within a gall, and the number of emigrants produced during the galling phase [18]. By examining these and other ecological, demographic and phenological differences (with special reference to measures of mortality, caste composition, and timing of gall opening), our aim is to elucidate the key selective factors that have influenced the evolution of sociality in the galling aphids and other fortress-defending organisms.
Ecology of P. populi
The average gall volume at the end of P. populi's galling phase was 1099 ± 458 mm3. Larger galls tended to contain greater aphid populations, increasing by approximately 1 aphid for every 70 mm3 of gall volume (R2 = 0.31, F1,19 = 8.62, P = 0.008). Correspondingly, larger galls also tended to allow the emergence of increased numbers of alate fundatrigeniae (with the number of these emigrants estimated to increase by approximately 1 for every 50 mm3 of gall volume, R2 = 0.21, F1,25 = 6.972, P = 0.014).
The index of developmental synchronisation in the gall of P. populi was calculated for all the destructive samples because the full range of instars was never in the gall at the same time. There was no significant difference between the resulting indices for each sample (F10,40 = 0.69, P = 0.72). The index calculated from the pooled samples indicated a high level of developmental synchronisation: on average, 0.92 ± 0.02 of the individuals in a gall was belonged within the same single instar.
The average number of fundatrigeniae in destructively-sampled galls was 42.9 ± 20.3. (The fundatrigeniae were the only morph in the gall other than the foundress.) This value accorded well with the average number of alate fundatrigeniae that left the monitored gall and were captured in bags: 41.0 ± 21.2. The density of aphids in galls of P. populi at the end of the season is therefore about 0.03 aphids mm-3.
The opening of the galls of P. populi is highly unusual. Wide fissures open in the gall starting at its apex and often extend so far that the gall's inner surface is turned entirely outward. A closed and an open gall are depicted in Table 2. Alate fundatrigeniae (which comprised virtually the entire aphid population) emerged immediately that the galls opened. This emergence was synchronised: the fundatrigeniae of all the bagged galls emerged on the same day, June 10 (see Figure 1A).
Table 2 Differences among P. populi, P. bursarius and P. spyrothecae in key defence-related components of their life history strategies
Comparison of the phenological characteristics of the three species P. populi , P. bursarius and P spyrothecae. (A) the emergence patterns, (B) the patterns of gall opening, and (C) the patterns of gall mortality (death of all the individuals within the gall). Splines are fitted to the data of P. populi and P. bursarius and regression curves are plotted for the combined data for P. spyrothecae. The data from the different field sites from which P. spyrothecae was sampled (CW: Cement Works, LP: Leys Playing Field, PS: Pembroke Sportsground, CF: Coton Footpath) are identified by the unique triangular symbols indicated in the legend.
Immediately following emigration from the gall of the fundatrigeniae (June 11), 42.0% of galls contained predators, 47.3% contained inquiline aphids, and 22.7% contained aphidopathogenic fungus. 87.3 % of the monitored galls (131 out of 150) were thus affected by phenomena that would have been fatal to aphids had they continued to inhabit them. This extremely high rate of gall mortality is shown in Figure 1C.
Ecology of P. bursarius
The average volume of P. bursarius galls at the time of peak population (June 20) was 521.0 ± 45.7 mm3. Galls with larger volumes again tended to contain larger numbers of aphids, with the aphid population increasing by approximately 1 for every 9 mm3 increase in volume (R2 = 0.45, F1,17 = 13.78, P < 0.001). The trend for a greater number of alate fundatrigeniae to emerge from larger galls was not significant (F1,28 = 3.39, P = 0.08).
As the average number of aphids in galls at peak population was 57.9 ± 10.9, the density of aphids in the galls of P. bursarius was approximately 0.11 aphids mm-3. The fourth-instar predominated at the time of peak population. The index value for synchronisation of aphid development was 0.67 ± 0.02. The galls were empty by July9, approximately 2.5 months after they had been formed (see Figure 2A).
Demographic composition of destructively-sampled galls. (A) P. bursarius: breakdown of aphid instars (mean number per gall ± SE) over the duration of the galling season; (B) P. spyrothecae: breakdown of aphid morphs/instars (mean number per gall ± SE) at the Cement Works field site over the duration of the galling season. Note that the y-axes are on different scales.
The galls of P. bursarius were open for most of the galling phase. Galls were closed for an initial 2–4 weeks but virtually all had opened by mid-June (Figure 1B). The approximately circular openings measured 1–1.5 mm in diameter. Emergence of alate fundatrigeniae began in mid-June and had concluded by the end of July (Figure 1A). Peak emergence was in early July. The mean number of fundatrigeniae emerging from each gall was 59.4 ± 8.8 for the entire season.
The rate of gall mortality was approximately constant over the entire galling phase (Figure 1C). The increase in mortality in response to opening of the galls was not especially pronounced because opening occurred very near to the beginning of the galling phase when failures (unrelated to predation) of the small founding populations were at their highest.
Ecology of P. spyrothecae
The total number of resident aphids within a gall correlated positively with gall volume in all cases. For example, at Pembroke Sportsground for the galls collected over the sampling period encompassing 14 August, 26 August and 8 September, 1999, the aphid population tended to be larger by one for every 3.5 mm3 increase in gall volume (R2 = 0.24, F1,131 = 40.36, P < 0.001). A positive correlation with volume was also true for the number of sexuparae emerging from the galls, which tended to increase by one for every 5 mm3 increase in gall volume (pooled result for all four sites: R2 = 0.33, F1,74 = 35.96, P < 0.001).
The aphid population peak occurred in mid-August. Pooling the two peak-population sampling events for every site, the average number of aphids per gall is 222.5 ± 6.4. The average gall volume at this time is 636.7 ± 13.4 mm3. The peak density of aphids inside the gall is therefore around 0.35 aphids mm-3. Soldiers and mature sexuparae were particularly numerous at the time of population peak and the peak in population co-incided with the opening of the galls. Fluctuations in the numbers of both virginoparous and sexuparous morphs over the entire field season are presented for each site in Figure 2B.
Aphid population size was influenced by the same factors that affected gall volume. Analysis of variance using the statistical model
Number of aphids = day + day × height + day × site(tree) + day × height × site(tree) + ε
showed that the first order interaction of day with site was highly significant (F3,1504 = 27.76, P < 0.001), just as population differences among trees remained detectable after having accounted for time-dependent and site-dependent differences (F4,1504 = 13.92, P < 0.001). Again, the height from which the galls were collected was not found to have a significant effect (P > 0.1 in both cases).
There was no relationship between the proportion of soldiers in a mature colony and colony population size at any site. Similarly, there was no relationship between the proportion of soldiers in a mature colony and gall size.
For P. spyrothecae, the index of developmental synchronisation was calculated at the time when all morphs were well-represented in the gall (August). First-instar soldiers predominated, making up 0.43 ± 0.01 of the gall. Death of a colony's foundress usually occurred 2–4 months after the gall's initiation. In mid-August (Day 90), approximately half of the galls contained a living foundress but all foundresses were dead by mid September (Day120).
Adult sexuparae began to emerge from the galls immediately that the openings were formed (see Figures 1A and 1B). The elliptical openings ranged from 1–2 mm in length (measured along the axis of the gall spiral) and 1–1.5 mm in width. Emergence continued for a period of almost three months. Peak emergence occurred in the month of September at the middle of the emergence period. The average number of sexuparae that emerged from each gall over the course of the season is 128 ± 8.5. Gall mortality (i.e. when all the aphids died) increased markedly following the opening of the galls, with the rate of gall mortality at least doubling in all cases (Figure 1C).
All four phylogenetic techniques produced the same topology which is given in Figure 3. Maximum likelihood bootstrapping analysis gives this tree topology strong support. Bayesian analysis estimated the posterior probability of this topology to be 0.82. The tree with the next greatest posterior probability of 0.12 differed in just one respect: the positions of P. populitransversus and P. populi were exchanged.
Phylogram of nine species of Pemphigus. The three species that were subject to detailed ecological study are shown in bold typeface. Eriosoma ulmi is the outgroup. A parenthetical "S" indicates that the species possesses soldiers and, where underlined, highly aggressive soldiers with specialised defensive morphology. The absence of soldiers is denoted by a parenthetical "N". Arrow (A) indicates the likely origin of soldiers, arrow (B) indicates the evolution of highly aggressive soldiers, and arrow (C) indicates the evolution of highly aggressive soldiers along with loss of host-alternation and the gain of dimorphic nymphs. The numbers on the branches give the number of times out of 100 that a bootstrap replicate recovered the branch, and the scale indicates units of 0.01 expected substitutions per nucleotide position.
Bayesian inference was also used to test support for monophyly of species known to possess soldiers. In the unconstrained analysis just described, if we assume that P. trehernei and P. populitransversus both possess soldiers, the cumulative posterior probability of a phylogeny containing a clade which is monophyletic for presence of soldiers is 0.95. On the other hand, the same analysis estimates that there is zero probability of a tree with soldier monophyly if we assume that neither P. trehernei nor P. populitransversus possesses soldiers. A second analysis which constrained P. gairi, P. bursarius, P. obesinymphae, P. protospriae, and P. spyrothecae to be monophyletic was also conducted. The difference in the harmonic means of the likelihood values from each of the two analyses was used to estimate the Bayes factor of 27.88. This value provides strong evidence that the model with the constraint of a five-species monophyletic soldier clade should not be accepted over the unconstrained model [19]. However, the difference resulting from comparison of the unconstrained model to a constrained model which adds P. trehernei as the sixth species of the monophyletic clade is less marked, as indicated by the considerably weaker Bayes factor of 12.48.
Life-history and ecological traits
Detailed ecological investigation of the three study species provided a clear demonstration of which factors are likely to have had the predominant influence in producing divergent defensive strategies through evolutionary optimisation of the aphids' life histories (Table 2).
Duration of the galling phase
The duration of the galling phase was approximately two months in P. populi, approximately three months in P. bursarius, and between five and six months in P. spyrothecae. P. populi is thus exposed to the selective environment of the primary host for the shortest time, while P. spyrothecae is exposed for the longest.
Aphid density
The lowest aphid density occurred in the galls of P. populi and was up to four times greater in P. bursarius galls and up to twelve times greater in P. spyrothecae galls. These density figures also reflect interspecific differences in the size of aphid populations supported in a gall. Whereas the P. bursarius peak populations (57.9 ± 10.9) were only slightly bigger than those of P. populi (42.9 ± 20.3), the peak populations of P. spyrothecae were more than four times the size (222.5 ± 6.4). Population size and density are likely to relate to factors such as the duration of the galling phase and the robustness of a colony following negative influences such as predation. Population size and density are also likely to have an integral influence on gall hygiene. It was noted that detritus such as moulted cuticle and waste such as wax-covered honeydew accumulated in the bottom of galls of P. populi. This was not the case for galls of P. bursarius and P. spyrothecae, the inhabitants of which push detritus and waste out of the holes in their galls [8, 9].
Gall opening
The opening of the gall saw a rise in aphid mortality in all three of the species. This opening releases a colony's winged migrants, but it also compromises the integrity of the gall as a physical boundary to predation [11] and, in P. spyrothecae, is the reference point for the specialised spatial distributions of the various gall morphs [20]. The potential risk of predation seems especially high for P. populi, the species without soldiers. Although alate fundatrigeniae of this species departed from the gall immediately that it opened, almost 90% of the galls suffered from potentially disastrous incursions by predators, inquilines and pathogens on the day of opening. Whereas the opening of galls of P. populi takes the form of extensive rupturing of the gall, opening in P. spyrothecae and P. bursarius is a more subtle process which involves the formation of a small hole (<2 mm in diameter) in the otherwise sealed gall. The presence and size of these openings, is known to be a crucial factor which the aphids closely regulate and, if necessary, repair to minimise predation [11].
Alate emergence
Each of the three species had a unique time for emergence, and there was little overlap among the emergence periods. The emergence of P. populi occurs first, in early June, and is followed by P. bursarius in June-July and finally by P. spyrothecae in August-October. There is also a striking difference between the durations of the emergence periods. P. populi emerge in a single pulse, with all the winged fundatrigeniae leaving their galls in virtually one day. The fundatrigeniae of P. bursarius emerge over a period of up to two months, while emergence of the sexuparae of P. spyrothecae can continue for more than three months (Figure 1).
Synchronisation of development
The rapid emergence of P. populi in particular, but also of P. bursarius, is facilitated by a high degree of developmental synchronisation. More than 90% of the aphids within a gall of P. populi tended to be of the same instar and up to 60% were of the same instar in galls of P. bursarius. Being from host-alternating species, these instars were soon destined to migrate from the gall. In contrast, first-instar soldiers (none of which are destined to emerge) comprised the greatest single morph in galls of P. spyrothecae. Despite being the most prevalent morph, the concurrent presence of a number of other morphs meant that these soldiers accounted only for approximately 40% of aphids.
Phylogenetic inferences
The three species that were subject to detailed ecological study are spread across the partial phylogenetic tree presented in Figure 3. The likely origin for the evolution of defensive behaviour is at the branch (point A) which gives rise to the six uppermost species of the tree. Behavioural defence is known for five of these six species, with P. trehernei representing the single exception. The loss of defensive behaviour on the P. trehernei branch must remain putative until thorough investigation is made but, if confirmed, the loss is likely to be associated with a life-history shift away from the primary host: the species has not been found to form galls in natural conditions [21]. Although the partial phylogeny suggests just one origin and one loss of the basic defence trait, it also suggests that pronounced defence through highly aggressive soldiers has evolved twice, once in the case of P. obesinymphae and once in the case of P. spyrothecae (points B and C on Figure 3). In the latter species, this increased defence is likely to have been supported by the evolution of dimorphism and the loss of all secondary hosts. The phylogeny certainly supports the hypothesis that P. spyrothecae evolved from a host-alternating ancestor [22].
Although detailed consideration of the evolution of gall forms (beyond their defensive implications) is outside of the scope of the current work, we note that the gall morphologies described by Blackman and Eastop [17] correspond well with the groups borne out in our molecular phylogeny. There is evidence of phylogenetic conservation of gall morphology in the facts that P. gairi has an elongate midrib gall, P. bursarius, P. obesinymphae and P. populitransversus have round petiole galls, while spiral petiole galls occur only in the P. protospirae/P. spyrothecae subgroup which is therefore probably monophyletic. The apparent phylogenetic conservation of gall morphology may also indicate the likely position of several species that are currently absent from our tree. For example, species such as P. matsamurai have galls that, like those of the study species P. microsetosus and P. populi, form on the leaf and dehisce in a secondary position to the original opening. On the other hand, with the possible exception of P. trehernei (which apparently has a unique biology and forms galls only rarely), our phylogeny does not include the various species of Pemphigus that form twig galls. These species are likely to belong to a separate subgroup, the position of which is yet to be elucidated. Intriguingly, because this subgroup would include P. dorocola, a species known to have soldiers [23], a second independent origin of defensive behaviour would be required.
The species examined in our phylogeny represent merely a small proportion of the large number of Pemphigus species and those of other related genera. Monoecy and defence by soldiers is certainly known to occur in a number of species that were not included in the current research. Incorporating these species into future studies will facilitate the expansion and, possibly, the amendment of the evolutionary relationships that we have inferred. Broader phylogenetic knowledge combined with improved behavioural and ecological knowledge will undoubtedly provide answers to some of the most interesting questions regarding the relationship between the selective environment and trends in social evolution.
Life-history strategies
Hide and run
P. populi has a short galling phase. The short time in the gall means that the time exposed to the predators of the primary host is reduced. Maintaining a completely sealed gall further isolates the colony from external influences. The build-up of waste is not a problem because colonies are relatively small and sparsely populated. Because the closed gall provides an extremely effective barrier to predation, soldiers are not required. The only time that the colonies are susceptible to predation is during the emergence period. This period is kept extremely brief so that predation is minimised. The pulse of emergence is facilitated by synchronising the development of the instars so that all are mature alate fundatrigeniae at the time of gall opening.
Stay and fight
P. spyrothecae has a long galling phase and a long emergence period. These extended periods can be sustained because the species possesses morphologically specialised, aggressive soldiers that are able to defend their colonies against predation. The long time in the gall under the protection of soldiers means that it is possible to increase a colony's reproductive success by having multiple generations and staggered development during the galling phase. The colony size is larger than those of the other species and aphids of P. spyrothecae live at a high density within their galls. The problem of the accumulation of waste at such high densities is controlled by having an opening to the gall by which the waste can be removed. The opening of the gall is relatively small and can be effectively guarded by the soldiers. A sustainable trade-off between preventing predators from entering and allowing migrants and waste to leave is thus possible.
A mixed strategy
P. bursarius has a galling phase and emergence period of moderate length. These periods of exposure can be borne by having moderately aggressive behavioural, but not morphological, soldiers. Galls have a small ostiole for most of the galling period. As is the case for P. spyrothecae, the presence of these ostioles can be supported because a balance can be found between ejecting waste and detritus from the gall and repelling predators with soldiers. The moderately high density of aphids makes this waste removal advantageous. Prolongation of the emergence period is prevented because the development of the immature aphids, which are all offspring of the foundress, is largely synchronised.
Key correlates of sociality
A variety of factors likely to be significant in the evolution of aphid soldiers have been identified. Increased defence investment is correlated with increasing aphid density, longer galling periods, longer periods of alate departure, small gall openings, and decreasing developmental synchrony. It is certain that predation pressure, an understudied factor, will have the central role in the evolution of defence. This assertion has been borne out, albeit in somewhat general terms, by the empirical (e.g. [11, 24, 25]) and theoretical studies that have dealt with the issue to date (e.g. [26–30]).
Another pivotal factor influencing the degree of defensive specialisation is whether loss of host alternation or other factors have influenced each species' innate predisposition for producing dimorphic offspring. Because dimorphism can facilitate the evolution of specialised defenders, it may well also result in increased fitness. Particularly in the genus Pemphigus, the loss of host alternation is likely to be a key determinate of this predisposition to dimorphism [22]. P. spyrothecae is the only study species that is not host-alternating and the loss of host alternation is likely to have increased the length of its galling phase. Certainly, other species that are restricted to the primary host, such as P. monophagus [31], also spend a relatively long period in the gall and have dimorphic first instars.
We hope that, by way of example, the current study has succeeded in exposing the excellent prospects that the genus Pemphigus holds as a model clade that can advance our understanding of the non-genetic influences upon the continuum that is sociality.
Study species
Typical Pemphigus life cycle
The majority of Pemphigus species have host-alternating life cycles in which periods spent on the woody primary host (poplar trees; Populus nigra in all three species described below) are succeeded by periods spent on herbaceous secondary host plants [17]. In spring, the founding female hatches from an egg which overwintered on the bark of the primary host and this foundress initiates a gall on the poplar leaf which is species-specific in its architecture. The foundress parthenogenetically produces the gall generation of tens to hundreds of individuals which, when mature, are specialised winged morphs (called fundatrigeniae/primary-host virginoparae) which migrate to the secondary hosts. Upon reaching the secondary host, the fundatrigeniae give rise (asexually) to wingless morphs (called virginoparae) which may dwell on the roots for a number of asexual generations before producing other winged migrant morphs (called sexuparae) which return to the primary host where they give rise to both male and female offspring. After mating, each female produces a single overwintering egg.
Pemphigus populi Courchet, 1879
The life cycle of P. populi adheres closely to the typical life cycle [32]. The monomorphic nymphs of P. populi exhibit no defensive behaviour [15], just as no other social trait has been found in this species.
Pemphigus bursarius (Linnaeus, 1758)
This species also has a life cycle that is broadly typical [33, 34]. However, the wingless secondary host females may occasionally produce a unique winged morph that can fly from one secondary host plant to another where it may found a new colony directly. The usual survival requirement of returning to primary host can thus be bypassed entirely [35].
Although the fundatrigenious nymphs of P. bursarius are monomorphic and without obvious morphologically specialised defenders, the first-instar individuals exhibit defensive behaviour. They respond aggressively to the cue of aphid haemolymph and actively attack potentially predacious syrphid larvae with their claws and stylets [15, 16].
The species has an atypical life cycle which is spent entirely on the primary host. Inside the gall, the foundress gives birth to a generation of aphids that are morphologically distinct soldiers in the first instar [36] but which nevertheless mature to wingless virginoparae that can produce both winged sexuparae (that migrate from the gall to the poplar bark) and more virginoparae [22, 24]. The virginoparous offspring of the second and subsequent galling generations are, once again, soldiers in their first instar. The life cycle described by Foster and Northcott [13] is correct except that we now know that these soldiers produced by the first generation of virginoparae are only facultatively sterile, and can mature to become reproducing virginoparae themselves if sufficient time remains before the leaves of the host plant are shed.
The soldiers of P. spyrothecae are among the most aggressive in the genus [15]. They are heavily sclerotised, have enlarged legs and are able to kill predators by puncturing cuticle with their stylets and especially curved claws [24, 37]. The presence of soldiers is highly effective in reducing predation on their colony [25].
Ecological sampling
Destructive sampling
Galls were harvested from heights between 1 and 15 m above ground level. These galls were immediately placed in separate sealed vials and taken to the laboratory within 6 h of collection. Each gall was measured to provide for volume calculations and examined for the presence of natural openings or external damage before it was broken open. Aphids were removed from broken galls with a soft brush and scored by instar and morph under a dissecting microscope. An index used for all three species to measure the synchronisation of development of aphids within their galls was created by taking the average proportion of the gall population represented by the predominant morph/instar.
Non-destructive monitoring
Phenology of the gall
Randomly-selected galls between 1 and 15 m above ground level were marked and numbered with brightly-coloured polyamide tape. At intervals specified below, each gall was externally examined for damage and the presence of natural openings. The dimensions of the gall were taken with a Vernier calliper (accurate to 0.5 mm) to track changes in volume. The apparent condition and quantity of the aphids within the gall was also estimated. Although the precise condition could only be obtained by destructive sampling, it was possible to discern if the aphids were dead or if the gall was empty.
Emergence of alates
The galls of a randomly-selected subset of those marked for monitoring were bagged securely with the feet of sheer, white, polyamide stockings. The alates that accumulated in these bags were periodically transferred to sealed polyethylene bags to be counted in the laboratory.
Emergence of alates provides a direct measure of the reproductive success of the galling phase of a clone. In P. spyrothecae, each sexupara that emerges from the gall will, if it survives, produce two males and between two and six females, and can thus provide for up to 6 overwintering eggs via which its genes can prevail [38]. In P. populi and P. bursarius, each fundatrigenia that leaves the gall can found an asexual secondary-host population [32, 33].
Sampling of P. populi
Field site
P. populi was sampled from a field site (43°34'28" N, 3°49'40" E) in the village of Saint Jean de Védas, France. This small area of parkland contained 30 black and Lombardy poplars.
P. populi was sampled between 1 May and 30 June, 2000. We destructively sampled 20 galls from up to 5 trees three times during this period. 150 galls (75 galls on each of 2 trees) were monitored; and 50 of these (25 galls on each tree) were bagged to gauge emergence of fundatrigeniae.
Sampling of P. bursarius
The site (52° 11' 2" N, 0° 10' 16" E) near Cambridge from which P. bursarius was sampled is a small reserve dominated by a single large Lombardy poplar surrounded by several smaller conspecifics.
P. bursarius was sampled from 28 May-26 July, 2001 at 1–2 week intervals. 30 galls were destructively sampled on seven occasions (a total of 210 galls). 70 galls were marked for monitoring and 50 of these were bagged to measure emergence of fundatrigeniae.
Sampling of P. spyrothecae
P. spyrothecae was sampled at four field sites in the vicinity of Cambridge, UK. The site 'Coton Footpath' (52° 12' 31" N, 0° 4' 8" E) is a playing field and reserve that contained 10 mature Lombardy poplars (Populus nigra var. italica). The site 'Cement Works' (52° 11' 49" N, 0° 9' 52" E) is an area of parkland on which more than 50 mature Lombardy poplars grew close together. The 'Leys Playing Field' (52° 11' 22" N, 0° 6' 58" E) is a sports field that contained 16 mature Lombardy poplars. 'Pembroke Sportsground' (52° 11' 25" N, 0° 6' 5" E), also a sports field, contained 20 mature Lombardy poplars.
P. spyrothecae was sampled between 11 May and 22 October, 1999. These dates were used to set the reference frame for interspecific comparisons. Thus, May 11 is set as Day1 and October 22 is set as Day165 for all three species. Destructive sampling and monitoring were conducted simultaneously, at 2–3 week intervals. At each sampling event, 50 galls were destructively sampled: 10 galls (5 from a height of 1 m and 5 from 10 m) were harvested from each of 5 randomly-selected trees. As all the field sites were sampled 9 times, 1800 galls were collected over the entire field season. Fifty marked galls were monitored galls on each of 2 trees at each field site from the beginning of the season until all the aphid colonies under observation had died. Emergence of sexuparae was recorded on a subset of 20 of the 50 monitored galls on each tree. Thus 400 galls were monitored in total, 100 galls at each field site. 40 galls were bagged to measure alate emergence at each site, giving 160 bagged galls across all the sites.
Construction of phylogeny
Aphids from eight species of Pemphigus (P. bursarius, P. gairi, P. obesinymphae, P. populi, P. populitransversus, P. protospirae, P. spyrothecae and P. trehernei) were collected and preserved in 95% ethanol. Individual alate aphids were freeze dried and genomic DNA was extracted using standard procedures. An area spanning part of the mitochondrial Cytochrome Oxidase I and II genes was amplified under standard PCR conditions using primers mt2793+ (5'ATACCTCGACGTTATTATTCAGA) and mt3660- (5'CCACAAATTTCTGAACATTGACCA). These, and the following internal primers were used in subsequent re-amplification and sequencing: mt2993+ (5'CATTCATATTCAGAATTACC), mt3175+ (5'CATGAC/TCATACAATTTTTATTAT) and mt3339- (5'GGA/GGATTTAATTTCATCTATT). The internal primers were developed by Stern [39] for use in sequencing P. microsetosus and species from the related aphid family Hormaphididae. Primer names indicate position relative to the Drosophila yakuba mitochondrial sequence [40].
PCR fragments were visualised by running out on a 2% agarose gel, then purified by cutting the bands from the gel and extracting the DNA by centrifugation followed by a phenol/chloroform extraction. A fraction of each of the resulting samples was re-amplified with internal primers and ethanol precipitated to obtain DNA for sequencing. Automated DNA sequencing was carried out using the ABI Prism dye terminator cycle sequencing kit with AmpliTaq DNA polymerase FS (Perkin-Elmer) and the ABI 310 genetic analyser (Perkin-Elmer). Results were analyzed and sequences aligned by hand using Seqed v. 1.03 (Applied Biosystems), with further sequence editing in preparation for phylogenetic analysis done using MacClade v. 3.05 [41]. The sequence of P. microsetosus, which had previously been used in Stern's phylogenetic analysis [39] was obtained from the author and also included in the analysis.
766–845 b.p. were sequenced for each species. Length variation was included in subsequent analyses, with deletions considered as a "fifth base". Length variation consisted of a single base deletion at base 223 in P. bursarius, P. obesinymphae, P. microsetosus and Eriosoma ulmi, within the tRNA region. Where bases could not be resolved accurately, the standard IUPAC codes for base uncertainties were used. All ten sequences were submitted to the EMBL/GenBank/DDBJ nucleotide databases [EMBL:AM748711–AM748720].
Four different techniques were used to estimate the phylogeny: (1) maximum parsimony was implemented with the heuristic algorithm of the computer program PAUP v. 3.1.1 [42] using 10 random addition sequences and all other options as default; (2) neighbour joining was performed using the NEIGHBOR program of the PHYLIP v. 3.53 software suite [43] and maximum likelihood distances for the neighbour joining algorithm were calculated with the DNADIST program of PHYLIP with a transition/transversion ratio of 0.72 and four categories of substitution rate for the three codon positions (0.40, 0.16, and 0.56) and the tRNA region (0.07); (3) the maximum likelihood phylogeny was estimated with the program fastDNAml v. 1.1 [44] using a global search starting from the maximum parsimony tree and the transition/transversion ratio and categories of substitution rate used in the neighbour joining analysis; and (4) Bayesian inference of phylogeny was performed using MrBayes v. 3.1.2 [45]. The transition/transversion ratio was fixed at 0.7 but otherwise default options were accepted for priors (flat priors for the rate matrix and branch lengths, uniform gamma shape parameter, equal site-specific rates) and Markov-Chain Monte Carlo parameters (four concurrent Markov chains and two runs which, after full convergence, were sampled every 100th generation).
All trees were rooted by comparison with the predefined outgroup Eriosoma ulmi from the pemphigine tribe Eriosomatini.
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NP acknowledges the generous financial support of the Cambridge Trusts and the Royal Society.
Department of Zoology, University of Oxford, South Parks Road, Oxford, OX13PS, UK
Nathan Pike
Department of Zoology, University of Cambridge, Downing Street, Cambridge, CB23EJ, UK
John A Whitfield & William A Foster
John A Whitfield
William A Foster
Correspondence to Nathan Pike.
NP designed, conducted and analysed the ecological research, contributed to the phylogenetic analysis, and prepared the manuscript. JAW designed and conducted the molecular research, and contributed to the phylogenetic analysis and manuscript. WAF assisted in planning the research and contributed to the manuscript. All authors approved the manuscript.
Pike, N., Whitfield, J.A. & Foster, W.A. Ecological correlates of sociality in Pemphigus aphids, with a partial phylogeny of the genus. BMC Evol Biol 7, 185 (2007). https://doi.org/10.1186/1471-2148-7-185
Defensive Behaviour
Aphid Population
Primary Host


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text:
Helen Mar Kimball (August 22, 1828 – November 13, 1896) was one of 30 to 40 plural wives of Joseph Smith, founder of the Latter Day Saint movement. She was sealed in marriage to him when she was 14 years old. After his death when she was 16, she married Horace Whitney "for time"; Whitney was the brother of another of Smith's wives. She bore eleven children with Whitney, the first three of whom died at, or soon after, birth. Their son, Orson F. Whitney, became an apostle in the Church of Jesus Christ of Latter-day Saints (LDS Church).

Biography

Early life
Kimball was born in Mendon, New York, as the third of nine children born to Heber C. Kimball and Vilate Murray. She was the only daughter to survive, and grew up being very close to her younger brother, William. As the only daughter, she was somewhat pampered by her parents . Kimball was three years old when her parents were baptized into the Latter Day Saint church in 1832. Kimball's family moved from Mendon to the church headquarters at Kirtland, Ohio, in the fall of 1833. When her father was called to be a church apostle in 1835, he was required to travel on missions and be away from home for significant lengths of time .

Kimball was baptized by Brigham Young in the Chagrin River during the winter when the river was frozen over. In order for her to be baptized, her father had to cut a hole in the ice. Kimball later wrote that she was not bothered by the cold water because she had "longed for this privilege" and that she "felt no cold or inconvenience from it" .

In 1838, the Kimball family moved from Kirtland to Far West, Missouri, to join church members moving there. Their arrival in Far West occurred soon after the Battle of Crooked River, and tensions between the Mormons and Missouri residents were beginning to reach a peak.

In early 1839, the family was forced to leave Missouri as a result of the Extermination Order issued by Governor Lilburn Boggs. As they left during the middle of winter, Kimball remembered how they had to keep walking in order to avoid freezing . The family eventually arrived in the town of Commerce, Illinois, which later was renamed as the city of Nauvoo. Kimball's father eventually built a house in Nauvoo near the temple lot. Her father enjoyed rising importance within church leadership and became a very close associate of Smith.

Marriage to Joseph Smith

According to Kimball, her father wanted to improve his standing by making a link between his family and the family of Joseph Smith .

Scholar Todd Compton has described the reason for the marriage as follows:

The prophet's marriage to her seems to have been largely dynastic—a union arranged by Joseph and Heber to seal the Kimball family to a seer, church president, and presiding patriarchal figure of the dispensation of the fullness of times .

In the spring of 1843, when Helen was 14 years old, her father described the doctrine of plural marriage to her. He asked if she would consent to be "sealed to Joseph" . Helen described her reaction to this proposition:

My father was the first to introduce it to me, which had a similar effect to a sudden shock of a small earthquake. When he found (after the first outburst of displeasure for supposed injury) that I received it meekly, he took the first opportunity to introduce Sarah Ann [Whitney] to me as Joseph's wife" .

Smith gave Helen 24 hours to respond to this request. The girl consented only after Smith explained to her that it would ensure her eternal salvation, along with that of her family. Helen was sealed to Smith in May 1843 when she was 14 and he was 37. The marriage was kept secret, and Helen continued to live with her parents .

Initially, Helen despised the concept of polygamy, stating that, "seeing the trials of my mother, felt to rebel. I hated polygamy with my heart." Later in her life, however, she became a vigorous defender of the practice and wrote a number of publications praising it (; ). With regard to her feelings about Smith's implementation of the practice, Kimball wrote,

It was a strange doctrine, and very dangerous too, to be introduced at such a time, when in the midst of the greatest trouble Joseph had ever encountered. The Missourians and Illinoisans were ready and determined to destroy him. They could but take his life, and that he considered a small thing when compared with the eternal punishment which he was doomed to suffer if he did not teach and obey this principle. No earthly inducement could be held forth to the women who entered this order. It was to be a life sacrifice for the sake of an everlasting glory and exaltation .

During the time that Helen lived in Nauvoo, she and Sarah Ann Whitney, who was also one of Smith's plural wives, became very close friends. According to Helen, she and Sarah were like "the two halves of one soul." Sarah's brother, Horace Whitney, married Helen "for time" after Smith's death in 1844 ().

Marriage to Horace Whitney
After Smith was killed in 1844, Kimball was 16 and had formed a relationship with 22-year-old Horace Whitney, a brother of another of Smith's wives. After a period of courtship, the two decided to be "married for time" on February 3, 1846 . Shortly before the exodus from Nauvoo, in the Nauvoo Temple, Kimball was married to Whitney "for time" and again sealed to Smith (deceased) "for eternity," with Whitney standing in as proxy for Smith. The following day, Whitney was sealed to Elizabeth Sykes (deceased) for eternity, with Kimball standing in as proxy for Sykes (;).

The Whitneys began the journey across the plains during the exodus from Nauvoo. They reached Winter Quarters, Nebraska in June 1846. Nineteen-year-old Helen bore her first child in May 1847 while her husband was away on an expedition to the Salt Lake Valley. The child was stillborn . In August 1848, while on the plains during the journey west, Helen had another child, who died shortly after birth. This birth resulted in complications to Helen's health, which almost resulted in her death. Following a long battle to regain her health, Helen bore her third child, who was born and died in September 1849 . She eventually bore a total of eleven children with Whitney . They were married for 38 years before his death. Their son, Orson F. Whitney became an LDS Church apostle.

Helen became a journalist, writing articles for the Deseret News and the Woman's Exponent. She also wrote the pamphlets "Plural Marriage as Taught by the Prophet Joseph" and "Why We Practice Plural Marriage", which defended the truthfulness and uprightness of this practice.

In 1896, Helen Kimball Whitney died at the age of 68 in Salt Lake City, Utah.

Controversy regarding Helen's marriage to Joseph Smith

The marriage of Helen Mar Kimball and Joseph Smith has long been a subject of controversy, most often with regard to her age at the time of the marriage. Jon Krakauer, in his book Under the Banner of Heaven (2003), says that of the women married to Smith: "Several were still pubescent girls, such as fourteen-year-old Helen Mar Kimball" . During a 2003 interview, Krakauer said in 2003: "They will not like the fact that I point out that Joseph Smith told 14-year-old girls 'God says you should marry me, you and your family will be exalted to heaven.' His way of getting laid doesn't reflect well on him."

Responding to Krakauer's characterization of Kimball's marriage to Smith, Mormon author Craig Foster states,

Falling into the same trap as many people and even some historians, he places his own modern values onto another place and time and, when their marriage patterns do not conform to his worldview, he looks upon it and writes about it with an open-mouthed, suitably shocked, and offended approach .

Foster cited examples of women marrying at a young age in colonial America, arguing that it was a common practice at the time. .

Notes

References

Jeni Broberg Holzapfel and Richard N. Holzapfel, eds., A Woman's View: Helen Mar Whitney's Reminiscences of Early Church History. Provo: Religious Studies Center, BYU, 1997.

1828 births
1896 deaths
Latter Day Saints from New York (state)
Latter Day Saints from Ohio
Latter Day Saints from Illinois
Latter Day Saints from Utah
Burials at Salt Lake City Cemetery
Mormon pioneers
Mormonism and polygamy
People from Mendon, New York
American pamphleteers
American women journalists
Child marriage in the United States
Wives of Joseph Smith

redpajama_set_name: RedPajamaWikipedia


text:
How Branded Entertainment Is Holding Up Amid Recession
By Big Fuel Staff in Trends
10 years ago	0 0
Madison and Vine's Andrew Hampp recently spoke with Ogilvy Branded Entertainment honcho Doug Scott about branded entertainment's place in the new econonmy.
PQ Media curbed its projections for branded entertainment as early as last February, saying the category's growth would slow to the high single digits this year after increasing 13.9% to become a $25.4 billion business in 2008. Product placement and integrations accounted for 15% of that business in 2008, with the rest going to events, sponsorships and other branded projects.
However, according to Scott the category has solidified it's importance and continues to prove itself as a necessecity in the new media landscape.
Read the full article here:
http://adage.com/madisonandvine/article?article_id=134580
Q&A: Ogilvy's Doug Scott on His Favored Metrics and Why Single-Channel Beats Product Placement
by Andrew Hampp
NEW YORK (AdAge.com) — As marketers rein in spending on everything from national TV to the Super Bowl, whither branded entertainment?
Helping lead the growth in recent years is Doug Scott, head of Ogilvy & Mather's branded-content division. Since joining Ogilvy in 2006, Mr. Scott has guided a diverse client roster through all aspects of the entertainment industry, from scripted TV (Unilever's Pond's on USA's "The Starter Wife") to music (a just-launched campaign for Kraft's Crystal Light featuring a new song from R&B songstress Estelle) to cable news (IBM's "The Business Of Innovation" on CNBC).
Although product placement and integrations have been the fastest-growing sector of branded entertainment, Mr. Scott has been similarly focused on single-channel entertainment programs such as microsites, branded video content and custom-created cable TV sponsorships.
"The need for accountability in marketing expenditures is greater than it's ever been, and I think the way that we're approaching branded content is somewhat unique in that it's a holistic approach," he said.
A more narrowly targeted model also enables Ogilvy to track a program's direct affect on sales. The agency was able to show, for instance, that Hellman's Mayonnaise doubled its year-over-year sales growth after launching a microseries, "In Search of Real Food," on Yahoo Food in 2007. Last summer, Dupont paint saw a sales boost after sponsoring a rebuilding initiative of Greensburg, Kan., that aired across Discovery Communications networks such as Discovery Channel, TLC and Planet Green.
Mr. Scott is aiming for similar results with new efforts such as "Digital Cribs," a microsite recently launched to drive sales for Cisco's consumer-products line and Estelle's new single, "Star," recorded exclusively for a new Crystal Light campaign that premiered last Sunday during E! channel's pre-Grammy red-carpet coverage.
Madison & Vine recently spoke with Mr. Scott to find out where branded entertainment is prioritized in clients' budgets for 2009; what's happening with the evolving metrics model; and why product placement should no longer be the focus of advertisers' branded-entertainment strategies.
Madison & Vine: How is branded entertainment holding up at a time when advertisers are looking to cut any kind of budget regarded as experimental or off-strategy?
Doug Scott: Clients are definitely looking to include it in their overall marketing mix, and we're getting a lot more interest in entertainment initiatives than I've seen in the past. There's still experimentation taking place, where clients are dipping their toe in before they make larger investments … in entertainment marketing. But a larger part is due to the fact that brands are looking for alternatives to traditional measured media to find more effective ways to engage the audience in an environment where you have screen-shifting and time-shifting and viewer control.
M&V: Because of the heightened demand for incorporating entertainment into all parts of a brand's strategy, what metrics are you favoring these days?
Mr. Scott: The first metric for us obviously is sales. That sales metric is really driven by two other metrics, one of which is a brand metric: What was the brand awareness, the brand lift, the brand shift? How are you making that brand more culturally relevant in an environment where the engagement with brands is floating to an audience much greater than it's ever been?
Then there's the media metric. We firmly believe that branded content really establishes this idea of working content, which can deliver more reach or impressions than that of working media because of digital distribution channels. So if you have really good content that can live in a multiplatform environment, the consumer you can target within an overall audience will both passively view the content and actively engage with a brand, thus moving the consumer up on the brand's consideration line.
M&V: Some of your clients, such as Hellman's, are reinvesting as a direct result of sales impact. What steps are you taking at the creative level at Ogilvy to help measure that relationship for clients?
Mr. Scott: The nature and the approach of the programs themselves are all about the underlying ROI. We're not just looking at the entertainment itself in a silo. It's about how it can be incorporated into advertising, the PR strategy around that, promotionally how that can be activated at retail, how online can be used to engage the audience and subsequently qualify that audience into a subset of consumers. Then it's about: Where are the natural retail partnerships to extend from a [consumer-package-goods] standpoint? And, of course, distribution: How do we go mass, narrow-cast and single-cast? And across all that, recognizing there is an entertainment asset that can be extended in a lot of different ways.
M&V: Pond's was one of the biggest success stories for brand integration after "The Starter Wife" in 2007, yet you've been focusing a lot more on single-channel, narrowcasted programs such as "Digital Cribs" for Cisco lately. Are scripted integrations such as "Starter Wife" just more difficult to execute now, or are they proving to be less effective as they become more commonplace?
Mr. Scott: Product placement, to me, is an easy-to-have, not a must-have, especially in the context of a story. It's a very crowded marketplace to be able to cut through, to be able to resonate when the receiver of that message is entertainment. If you really look back into the early days of television, the soap opera was an exchange the brand was making with the audience, in this case the housewives who were watching as they ironed their husbands' shirts. In return the brand got their time or attention, and in doing so began some kind of brand relationship, so the next time Mrs. Smith went to the market, she was more prone to buy P&G because she felt value exchange there.
I think we as marketers have lost that exchange because of the amount of things vying for our time — and extent of the disruption we all experience on a daily basis. When you get into a taxicab today, the minute you tell the driver where you're going and he starts the meter, the TV goes on, and you have to take a second out of your time and, in essence, take a second out of the conversation you're in to press "off." We live in a world of disruptive media. What value do I get from people infringing upon my space with their message?
M&V: So which networks are doing a good job managing that disruption?
Mr. Scott: NBC has been the most progressive network out there, in large part to [NBC Entertainment/Universal Media Studios co-chairmen Ben] Silverman and [Marc] Graboff in understanding what's old is new. The distribution platforms they have out there, with Hulu and a few other NBC digital channels, really give you the single-cast environment to take advantage of that. What that model does is it exhibits the changing economics of the business — and really assists the network to offset some of its risk with these programs, and develop unique marketing partnerships from the ground up where the brand really gets greater value than just 30-second spots.
Advertising Creative Innovation & Technology Marketing Media test Video

redpajama_set_name: RedPajamaCommonCrawl


Tell the Minor Hill TN you found them on ILocalFlorist.com!
ordering directly from local florist shops in Minor Hill Tennessee!
using an existing Minor Hill Tennessee address!
This directory was created to help you find a local florist shop in Minor Hill TN. Use the local florist shop's zip code for a list of Minor Hill Tennessee florists, flowershops and floral directory. The links to the local florist shops web sites in Minor Hill Tennessee are being provided as a service. By clicking on a link, you will leave the ILocalFlorist.com web site. The ILocalFlorist.com link to local florist shops web site in Minor Hill TN does not necessarily imply an endorsement by ILocalFlorist.com of any product, resource, or service on that local florist shops web site. ILocalFlorist.com cannot guarantee the content of the linked web sites.


text:
Getting to theroute of the problem
By wpengine	 1st June 2009 No Comments
Don't believe everything you read - an adage normally reserved for the susceptible readers of red tops and celebrity rags, but the same advice, it seems, should also be adhered to by truck drivers if they want to avoid (quite literally) being led up the g
Don't believe everything you read – an adage normally reserved for the susceptible readers of red tops and celebrity rags, but the same advice, it seems, should also be adhered to by truck drivers if they want to avoid (quite literally) being led up the garden path.
There have been a number of stories about HGV drivers getting stuck down narrow country lanes or up steep hills after following the instructions on their sat nav a little too stringently, causing untold damage to roads, countryside, surrounding buildings and the vehicle itself, not to mention disrupting residents and blocking access for other road users.
The problem has been attributed in part to foreign drivers not understanding instructions (so much so in some areas that pictorial signs have been erected warning drivers to ignore their sat navs) as well as the fact that manufacturers have so far focused primarily on the car market.
To help combat the problem, mapping agency Ordnance Survey has released "steep road" information to help sat nav providers route truck drivers more appropriately. The new data complements the existing information it published in 2006 on road weight, width and height limits, which it hopes will encourage sat nav manufacturers to release more products designed specifically for HGVs, helping to reduce the number of vehicles being sent down unsuitable roads.
Narrow lanes
Garmin has just launched the nüvi 465TF, a portable navigation device designed expressly to stop truck drivers from getting stuck down narrow village lanes and under low bridges. The system takes into account the vehicle's dimensions and load restrictions before adjusting the route to best suit the truck's profile using the available map data.
TomTom Work, the business subsidiary of TomTom, launched a navigation system for truck drivers at the end of 2008. More recently the company released TomTom GO 7000, which delivers real-time traffic information for commercial vehicle drivers, as well as reports about delays and alternative routes. At the same time TomTom Work released its WebFleet Dashboard, a reporting tool within the online fleet management application, TomTom WebFleet, which shows all analyses and trends in a single overview.
Thomas Schmidt, managing director of TomTom Work, says: "While the new TomTom GO 7000 makes professional navigation even more time and fuel efficient, the TomTom WebFleet Dashboard supports the management of fleets. Especially in times of economic difficulty, when costs have to be reduced and processes optimised, TomTom WebFleet Dashboard enables decisions to be made quickly, based on information that is displayed clearly and that can be quickly retrieved."
Whether it's improving efficiency, boosting productivity, tightening costs or optimising staff output, squeezing that little bit more out of what you have has never been so important.
Routeing software can help boost efficiency and create huge reductions in outgoings, says Optrak's Tim Pigden. "Common vendor literature says savings of between five and 15 per cent can be achieved in transport and transport management costs, which is typically achieved by reducing the number of vehicles used and reducing the time spent planning. However, there is still a huge dichotomy between people who use routeing software and have seen proven benefits, next to those who don't understand it, or don't want to understand it, and therefore don't use it."
Fuchs Lubricants was able to reduce the time it spent planning by more than 85 per cent after investing in the Optrak vehicle routeing package. Richard Halhead, Fuchs' managing director, says in addition, "we have improved all of our KPIs: on time or early delivery performance, cutting the pence per litre cost of each delivery, increasing the utilisation of vehicles and reducing mileage leading to a reduction in our carbon footprint."
The company is also less reliant on one member of staff with specialist knowledge, so is able to cope better when employees are absent, as other members of the team can still create effective loading plans and routes.
Better planning can also create huge savings financially. William Salter, managing director of Paragon Software Systems, says: "The first prerequisite of an efficient transport operation is to ensure it is planned effectively. The routeing and scheduling optimisation systems required for this are well established and continually advancing in line with hardware performance improvements and industry demands. This helps companies plan efficient schedules that maximise fleet utilisation and minimise operating costs."
After implementing Paragon's transport optimisation system furniture manufacturer Dams International is saving £750,000 annually, through better management of its drivers, fitters and 140-strong fleet.
To boost its offering, DHL's drinks logistics company Tradeteam chose to develop a transport management system in-house. Network manager Kevin Lawrence says: "By using the TMS, drivers have ease of mind when communicating with customers as they work with a clear summary of their daily activities and managers are able to drive efficiencies through knowledge gained from interpreting the information the system provides."
Tradeteam has seen around a 21 per cent rise in efficiency levels since implementing the system, which in turn "has allowed service improvements and enhancements to be funded – differentiating us from our competitors," says Lawrence.
When it comes to tackling inefficiencies Roger Marks, managing director of Aeromark, says a combination of technologies is best. "Routeing software helps managers and planners select the most effective routes. Scheduling applications ensure delivery sequences of delivery/collection locations are optimised. Sat nav helps to prevent drivers from wasting fuel by getting lost. Crucially, vehicle tracking shows where the driver has actually been and can help to analyse driver behaviour."
However, while a combination of technologies may well be the optimum solution, if a company is not in a position to spend huge amounts on multiple new systems Marks suggests "a 'must-have', without any doubt, is a reliable, robust, vehicle tracking solution, because if you can't measure you can't manage".
He says: "With the clear visibility of what is actually happening in the field, you can begin to ascertain where areas for potential improvement may lie in your business process. After the initial analysis, scheduling software can offer more efficient planning of deliveries and routes."
Conversely, Paul Palmer, chief executive of DPS, insists: "What good is knowing where a vehicle is if you don't know where it should be?" He reckons many companies get seduced by tracking as it is cheap, when in actual fact scheduling is more effective. "If you've got a good plan in place first then knowing where vehicles are is much more useful. You should begin by planning the most efficient route and then use tracking to help achieve that plan."
Before making any hasty decisions, Chris Greenwood, CEO of MapMechanics, advises companies to have a clear idea of what they are ultimately trying to achieve, because "while all technology has a part to play, not all technology will be able to help everyone," and making a mistake could end up being more costly. He adds that as a result of budgets being tighter, there has also been a move towards pay-as-you-go services.
A trend that hasn't gone unnoticed by DPS's Paul Palmer. He says: "There is still the traditional product which companies pay £15,000 – £20,000 for that we install and train up staff to use, but increasingly people don't want that. Now, customers in general would prefer to have it hosted on the internet as they don't have the cash flow to pay up front. Software as a Service is growing very fast.
"It is also particularly good for smaller companies, who traditionally couldn't afford to pay up-front licence costs, but find it more cost effective over the web as they don't need any other infrastructure. The web version of the software is taking routeing and scheduling into service areas that never thought they would need it."


text:
Q: How to create independent JPanel-Layers within JFrame and repaint each indiviually I'm searching for a way to paint several JComponents above each other (overlap) and still being able to individually access and alter them.
E.g. paint three JPanels with transparent backgrounds - each containing a circle, a rectangle or a line.
Afterwards, I'd like to change the appearance of the circle. The other two should not be repainted (similarly to layers in Photoshop).
My current project has a Jpanel with thousands of lines and I need to change a rectangle in the back on mouseover if I redraw the complete Jpanel each time it is very laggy.
Is there a decent way to accomplish that? Thank you already for your ideas!

A:
I need to change a rectangle in the back

You can invoke:
panel.repaint(rectangle); // or
panel.repaint(x, y, width, height);

to specify the Rectangular area to be repainted.

A: It worked quite fine - here is my code if anyone else has a similar problem!
the first image can be stored and displayed later on (buff)
Make sure to generate new BufferedImage (here canvas) when displaying again, as the transperency is lost otherways. Thanks to Gilbert Le Blanc
@Override
protected void paintComponent(Graphics g1) {
    //Create image:
    BufferedImage buff = new BufferedImage(mywidth, myheight, BufferedImage.TYPE_INT_ARGB);

    //write to image:
    Graphics2D g2 = (Graphics2D) buff.getGraphics();
    g2.setComposite(AlphaComposite.getInstance(AlphaComposite.SRC_OVER,(float) 0.01f));
    g2.setRenderingHint(RenderingHints.KEY_ANTIALIASING, RenderingHints.VALUE_ANTIALIAS_ON);
    g2.draw(xxxRectanglewhatever);

    //then - later draw image again
    BufferedImage canvas = new BufferedImage(mywidth, myheight, BufferedImage.TYPE_INT_ARGB);
    canvas.getGraphics().drawImage(buff, 0, 0, null);
    ((Graphics2D) g1).setRenderingHint(RenderingHints.KEY_ANTIALIASING,RenderingHints.VALUE_ANTIALIAS_ON);
    g1.drawImage(canvas, 0, 0, null);
    canvas.flush();
}

redpajama_set_name: RedPajamaStackExchange


This section is dedicated to Guttorm Vik, who wrote StrongED and StrongHelp, and to Fred Graute who is maintaining StrongED.
For my own convenience I have designed several StrongED modes, but as these have not been polished and properly documented I have kept them to myself. I will be making these available here in the future.
StrongHelp is such a useful way of giving help to a user that I try to provide a manual for my Software projects. I do not include this in the main download, so one way of getting a feel for my work is to get the manual first.
Having written an article for Archive about StrongED Search and Replace, I have made it into an A5 pamphlet, and followed this with another about Modes files. Comments and corrections are very welcome.
Search and Replace OvPro Search and Replace pamphlet.
Mode files OvPro Mode files pamphlet.


text:
Browsing by Composer "Lachner, Franz Paul"
Now showing result 2160 of 4276
Flute Concerto in D minor (Flute Concerto in D minor)
Lachner, Franz Paul ; Meier, Bruno; Prague Chamber Orchestra (1832) [15:58]
I. Allegro con brio (Symphony No. 1 in E flat major, Op. 32)
Lachner, Franz Paul ; Choo, Hoey; Singapore Symphony Orchestra (1828) [11:09]
I. Allegro moderato (Octet in B flat major)
Lachner, Franz Paul ; German Wind Soloists (1850) [10:01]
I. Andante - Allegro maestoso (Symphony No. 8 in G minor, Op. 100)
Lachner, Franz Paul ; Robinson, Paul; Slovak State Philharmonic Orchestra, Kosice (1851) [16:47]
I. Andante - Allegro (Symphony No. 5 in C minor, Op. 52, "Preis-Symphonie")
I. Andante maestoso (Septet in E flat major)
Lachner, Franz Paul ; Villa Musica Ensemble (1824) [11:07]
II. Adagio (Octet in B flat major)
II. Andante con moto (Symphony No. 5 in C minor, Op. 52, "Preis-Symphonie")
II. Andante (Symphony No. 1 in E flat major, Op. 32)
II. Andante (Symphony No. 8 in G minor, Op. 100)
III. Andante (Septet in E flat major)
III. Intermezzo (Suite No. 7 in D minor, Op. 190)
Lachner, Franz Paul ; Gunzenhauser, Stephen; Polish State Philharmonic Orchestra, Katowice (1881) [05:55]
III. Menuetto (Symphony No. 5 in C minor, Op. 52, "Preis-Symphonie")
III. Scherzo: Allegro assai (Symphony No. 1 in E flat major, Op. 32)
III. Scherzo (Octet in B flat major)
III. Scherzo (Symphony No. 8 in G minor, Op. 100)
III. Variationen und Marsch (Suite No. 1 in D minor, Op. 113)
III. Walzer (Ball-Suite in D major, Op. 170)
Lachner, Franz Paul ; Walter, Alfred; Slovak State Philharmonic Orchestra, Kosice (1874) [05:41]
II. Mazurka (Ball-Suite in D major, Op. 170)
II. Menuetto (Septet in E flat major)


An interview with Spanish guard Marc Garcia at the 2015 FIBA U19 World Championship in Crete.
-Smooth athlete. Very fluid and mobile.
-Good instincts as a scorer and shot-creator.
-Struggling badly with his jump-shot as of late. 29% for 3 in the ACB this season. 30% at this tournament. In a huge funk from beyond the arc in past year plus.
-Poor defender. Lacks toughness and strength. Intensity level leaves a lot to be desired. Doesn't close out with urgency. Doesn't fight through screens.
Outlook: Creative shooting guard with nice scoring instincts. In the midst of a difficult period, coming off a tough season in the ACB. Seems to have lost much of his swagger which made him such an intriguing player earlier in his career. Frame needs to improve substantially. Still flashes signs of intrigue with his ability to draw fouls, get in the passing lanes and make an occasional jumper, but needs to improve his defense, shooting consistency and become a much better finisher inside the arc to reach the tremendous potential he showed earlier in his career. Rights are owned by Barcelona, so will almost certainly be on loan again next year.
-What kind of defender can he become?
Outlook: Kind of a Spanish Nik Stauskas. Made a very strong impression in Mannheim. Appears to have grown, filled out physically, and improved considerably in the past 12 months. Clearly one of the best young prospects in Europe. Already producing at a very high level in the Spanish second division (LEB Gold), averaging 11 points in 24 minutes. Needs to continue to work on his body, keep improving defensively.
-Needs to improve passing skills and ability to create for others. Only dished out two assists in 231 minutes.
Outlook: Exceptionally talented young scorer. Looks like a baby but has a killer's mentality. Comes on the floor looking to put up points, but doesn't force the issue and plays within a team concept. Lack of strength affects everything he does at the moment, but he just turned 17 two months ago so he still has plenty of time to work on his frame.


text:
Because the app is modeled with the Twelve-Factor app architecture, all configs are stored in the environment.

When executing mix command, you should always make sure that the required system env are present.
You can use `source`, [nv](https://github.com/jcouture/nv) or a custom l33t bash script.

Every following steps assume you have this kind of system.

## Running the app

  1. Create your .env and .env.test config file.
  2. Install dependencies with `mix deps.get`.
  3. Create and migrate your database with `mix ecto.setup`
  4. Start Phoenix endpoint with `mix phx.server`

## Environment variables

The application needs the following environment variables:

```
PORT=
CANONICAL_HOST=

# Secret key. You can use `mix phx.gen.secret` to get one or use any secret string you want
SECRET_KEY_BASE=
SESSION_KEY=

# Url of the database
DATABASE_URL=
# Pool size of the DB connections. You may use 20 connections as a starting point.
DATABASE_POOL_SIZE=
```

It also supports these optional environment variables:

```
FORCE_SSL=
BASIC_AUTH_USERNAME=
BASIC_AUTH_PASSWORD=
```

## Linting

Use: `mix credo --strict`

redpajama_set_name: RedPajamaGithub


The Joint Research Centre (JRC) is the European Commission's in-house science service which employs scientists to carry out research in order to provide independent scientific advice and support to policies of the European Union.
In this catalogue, you can find an inventory of data produced by the JRC in accordance with the JRC Data Policy. The content is continuously updated and shall not be seen as a complete inventory of JRC data. Currently, the inventory describes only a small subset of JRC data.


text:
GM EV Mandate Announcement Draws Fire from All Sides
Posted By Alexandra Bly In Auto & Alternative Vehicles,Demand & Efficiency,Energy Politics,United States |
Full Story: E&E News @EENewsUpdates
Primary Author Maxine Joselow @maxinejoselow
Tennen-Gas/Wikimedia Commons
General Motors managed to alienate the full spectrum of potential allies with its call last week for the Trump administration to introduce an electric vehicle mandate, with other automakers complaining they'd received insufficient notice of the announcement, a fossil lobbyist accusing the company of looking for government handouts, and green groups accusing GM of greenwashing.
"If there was a way to make more people mad, I'm not actually sure I know what it is," an anonymous source told E&E News. "This seems like a stupid way to make literally everyone in Washington mad at you."
"General Motors Co. was hoping for a positive reaction when it made an eye-popping announcement [2] last week about boosting electric vehicles," E&E reports. "Instead, America's largest automaker was flooded with criticism."
Within the auto industry, sources told E&E that other companies found out about the release when GM circulated an embargoed news release a few hours before it went public. Its position broke with the Alliance of Automobile Manufacturers, which had just submitted comments to a review of Trump's proposed vehicle fuel efficiency rollback that were supposed to speak for all 12 member companies.
"We didn't know about it ahead of time," one source said. "I think their embargo lifted at 12:01. So we found out when everybody else did. I didn't even find out until seeing [GM CEO] Mary Barra's op-ed. We just were caught off guard like everybody."
Dan Becker, executive director of the Safe Climate Campaign, said other automakers might have supported GM's position if they'd known about it far enough in advance. And that missed opportunity had him skeptical about the company's intentions.
"When they really want something, they know how to go about trying to get it," he said. "But they just sort of shoved this out there, pissing everybody off." That action "gave me the sense that they really aren't serious about it. It was just for purposes of green-scamming, or pretending they're doing something environmental when they really aren't."
Fossil ally Thomas Pyle, president of the American Energy Alliance, said his group would stand against a GM proposal that he saw as a request for government handouts for electric vehicles. He also speculated the company might be counting on Trump departing the White House after the next U.S. federal election in 2020.
"GM's proposal is a tacit admission that they can't sell electric vehicles without government subsidies and mandates," Pyle told E&E. "Politically, it is a thumb in the eye" to Trump, with a proposal that "is surely dead on arrival in the White House".
Environmental groups cast the announcement as a bid to draw focus away from GM's support for the fuel economy rollback.
"With transportation accounting for more than one-third of our greenhouse gas emissions, the U.S. must continue its transition to clean, electric vehicles," said Gina Coplon-Newfield, director of the U.S. Sierra Club's Clean Transportation for All campaign. "However, General Motors' proposal is a desperate attempt to distract from their support for Donald Trump's efforts to roll back the clean car standards."
Luke Tonachel, director of clean vehicles and fuels at the Natural Resources Defense Council, cast the announcement in similar terms, calling it a "distraction from preserving the existing, commonsense tailpipe pollution standards."
URL to article: https://theenergymix.com/2018/11/05/gm-ev-mandate-announcement-draws-fire-from-all-sides/
[2] eye-popping announcement: https://theenergymix.com/2018/11/02/transport-canada-embraces-gms-ev-mandate-but-gm-canada-isnt-so-sure/


text:
Q: Exception java.lang.NoClassDefFoundError in Dynamic Web Application, Eclipse, JSF when I'm running my appication everything is ok to moment when I press the button and I'm redirecting to another xhtml page. Then eclipse console until I stop server display:
SEVERE: Servlet.service() for servlet [Faces Servlet] in context with path [/ChatAzure] threw exception [javax/servlet/jsp/jstl/core/Config] with root cause
java.lang.NoClassDefFoundError: javax/servlet/jsp/jstl/core/Config
    at com.sun.faces.application.view.JspViewHandlingStrategy.executePageToBuildView(JspViewHandlingStrategy.java:345)
    at com.sun.faces.application.view.JspViewHandlingStrategy.buildView(JspViewHandlingStrategy.java:154)
    at com.sun.faces.lifecycle.RenderResponsePhase.execute(RenderResponsePhase.java:100)
    at com.sun.faces.lifecycle.Phase.doPhase(Phase.java:101)
    at com.sun.faces.lifecycle.LifecycleImpl.render(LifecycleImpl.java:139)
    at javax.faces.webapp.FacesServlet.service(FacesServlet.java:594)
    at org.apache.catalina.core.ApplicationFilterChain.internalDoFilter(ApplicationFilterChain.java:305)
    at org.apache.catalina.core.ApplicationFilterChain.doFilter(ApplicationFilterChain.java:210)
    at org.apache.catalina.core.StandardWrapperValve.invoke(StandardWrapperValve.java:224)
    at org.apache.catalina.core.StandardContextValve.invoke(StandardContextValve.java:169)
    at org.apache.catalina.authenticator.AuthenticatorBase.invoke(AuthenticatorBase.java:472)
    at org.apache.catalina.core.StandardHostValve.invoke(StandardHostValve.java:168)
    at org.apache.catalina.valves.ErrorReportValve.invoke(ErrorReportValve.java:98)
    at org.apache.catalina.valves.AccessLogValve.invoke(AccessLogValve.java:928)
    at org.apache.catalina.core.StandardEngineValve.invoke(StandardEngineValve.java:118)
    at org.apache.catalina.connector.CoyoteAdapter.service(CoyoteAdapter.java:407)
    at org.apache.coyote.http11.AbstractHttp11Processor.process(AbstractHttp11Processor.java:987)
    at org.apache.coyote.AbstractProtocol$AbstractConnectionHandler.process(AbstractProtocol.java:539)
    at org.apache.tomcat.util.net.JIoEndpoint$SocketProcessor.run(JIoEndpoint.java:298)
    at java.util.concurrent.ThreadPoolExecutor$Worker.runTask(ThreadPoolExecutor.java:886)
    at java.util.concurrent.ThreadPoolExecutor$Worker.run(ThreadPoolExecutor.java:908)
    at java.lang.Thread.run(Thread.java:662)

I use:
eclipse, jsf2-1-7, jstl-1.2.jar, Tomcat v0.7.
I don't know what's wrong, I need help.

A: The mentioned class is part of JSTL. This is normally already provided by a full fledged Java EE container, but not in Tomcat as being a barebones JSP/Servlet container. You'd need to supply JSTL yourself along with the web application. It has ultimately to be placed in the webapp's runtime classpath.
You correctly mentioned jstl-1.2.jar (I assume that it's exactly the one as you could find in our JSTL wiki page), but this problem indicates that it has apparently not been placed at the right place. You need to place it in /WEB-INF/lib folder of the web project. This folder is covered by webapp's default runtime classpath. Nothing more needs to be done. If you have ever fiddled in the project's Build Path properties in an attempt to solve it, you should undo all those attempts.

Unrelated to the concrete problem: why are you still using legacy JSP? It has been deprecated and succeeded by Facelets (XHTML) since JSF 2.0. Make sure that you're reading up to date JSF 2.x resources and not JSF 1.x ones while learning JSF.


Dr. Lo was born in Washington and is excited about practicing family dentistry in his hometown. He strives to build long-term relationships based on dental health with his community. He seeks to provide a comfortable understanding of all the options and facilitate treatment that uniquely meets the needs of each person. He supports a minimally invasive approach to dentistry focused on prevention and the preservation of tooth structure whenever possible.
Dr. Lo enjoys active membership in the American Dental Associate, the Academy of General Dentistry, the Washington State Dental Association, and the Seattle-King County Dental Society. He has served on numerous councils and committees with the American Dental Society and is active in local leadership. He is active in National Dental Practice-Based Research with a Professional Research Assistant appointment by NYU and formerly an instructor at the Lake Washington Technical College in their Hygiene and Restorative Departments. He participates in many other study groups and continuing education courses which explore advances in the fields of restorative dentistry, implants, and periodontal therapy.
Dr. Lo is a proud 2002 graduate of the University of Washington Dental School. He grew up in the Bellevue/Redmond area of Washington, and attended Stevenson Elementary and Highland Middle School. He's also an alumni of Interlake High School on the Bellevue/Redmond border.
Outside his dental practice, Dr. Lo enjoys spending his free time with his family. he is happily married to his wife, Lisa, who is the mother of his two active sons, Ryan and Matthew. He loves getting outdoors in the scenic back country of Washington to fish, hike, and ski. He is also an avid sports enthusiast and closely follows the University of Washington Huskies and Seattle Seahawks.
Dr. Maurissa Kiefer is a California native who has fallen in love with all that the PNW offers. Dr. Kiefer received her D.D.S. degree from one of the nation's top dental schools, the University of the Pacific Arthur A Dugoni School of Dentistry, in San Francisco, CA.
Dr. Kiefer strives to provide her patients with comfortable, high quality care. Her philosophy is to provide comprehensive treatment to help each patient maintain their best oral health. Dr. Kiefer is dedicated to making sure that her patients feel comfortable and confident in the quality of their dental care.
Dr. Kiefer is a respected member of the American Dental Association, Washington State Dental Association, the Academy of General Dentistry, the Academy of Cosmetic Dentistry, and Seattle-King County Dental Society. Dr. Kiefer strives to expand her knowledge of the dental field by participating in numerous study groups as well as elite continuing education courses. Dr. Kiefer is also a certified Invisalign provider.
Outside of the office you can find Dr. Kiefer soaking up the sun while boating in the summer, skiing down the mountains in the winter months, and frequenting the gym to feed her fitness obsession.
Dr. Sengupta graduated from the University of Washington, School of Dentistry with Specialization in Periodontics and received a Master of Science Degree in Dentistry. She has extensive training in the field of Periodontics including Dental Implant surgery, Bone grafting, Periodontal esthetic surgery, and treatment of Advanced periodontal disease. Her professional training spans over 15 years of clinical and educational experience in various fields of Dentistry.
She is currently an Affiliate Associate Professor in the Department of Periodontics at the University of Washington, School of Dentistry. She is a member of the American Dental Association, Washington State Society of Periodontists, Washington State Dental Association and Seattle King County Dental Society. She is an active member of many study clubs and frequently presents lectures on clinical cases, techniques and innovations in Periodontics.
Dr. Sengupta's experience and expertise empowers her to drive a practice philosophy grounded in knowledge, skill and personal relationships with her patients. She treats her patients as family members with empathy and compassion and ensures a high level of ongoing periodontal health for them. She is friendly, cheerful, gentle, and compassionate with a pleasing chair-side manner.
Dr. Sengupta has been happily married for 10 years. She and her husband Tirthankar, have two children, Rohan and Riana. They live in Sammamish, Washington. In her free time, Dr. Sengupta enjoys baking, painting, traveling and spending time with her family.


The Apera Instruments ORP60-DA is the replacement ORP probe for PH60-Z, PH60S-Z, PH60F-Z, and PC60-Z. This probe features a Platinum ORP sensor with a double-junction structure. Designed for general water solution testing's ORP measurement.