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ORIGINAL RESEARCH article
Sec. Coral Reef Research
Volume 5 - 2018 | https://doi.org/10.3389/fmars.2018.00160
Interspecific Hybridization May Provide Novel Opportunities for Coral Reef Restoration
- 1Australian Institute of Marine Science, Townsville, QLD, Australia
- 2School of BioSciences, University of Melbourne, Melbourne, VIC, Australia
- 3School of Mathematics and Physics, University of Queensland, St Lucia, QLD, Australia
- 4Bio21 Institute, University of Melbourne, Melbourne, VIC, Australia
Climate change and other anthropogenic disturbances have created an era characterized by the inability of most ecosystems to maintain their original, pristine states, the Anthropocene. Investigating new and innovative strategies that may facilitate ecosystem restoration is thus becoming increasingly important, particularly for coral reefs around the globe which are deteriorating at an alarming rate. The Great Barrier Reef (GBR) lost half its coral cover between 1985 and 2012, and experienced back-to-back heat-induced mass bleaching events and high coral mortality in 2016 and 2017. Here we investigate the efficacy of interspecific hybridization as a tool to develop coral stock with enhanced climate resilience. We crossed two Acropora species pairs from the GBR and examined several phenotypic traits over 28 weeks of exposure to ambient and elevated temperature and pCO2. While elevated temperature and pCO2 conditions negatively affected size and survival of both purebreds and hybrids, higher survival and larger recruit size were observed in some of the hybrid offspring groups under both ambient and elevated conditions. Further, interspecific hybrids had high fertilization rates, normal embryonic development, and similar Symbiodinium uptake and photochemical efficiency as purebred offspring. While the fitness of these hybrids in the field and their reproductive and backcrossing potential remain to be investigated, current findings provide proof-of-concept that interspecific hybridization may produce genotypes with enhanced climate resilience, and has the potential to increase the success of coral reef restoration initiatives.
The rapid increase in atmospheric CO2 to levels not documented for millions of years (Hönisch et al., 2012) and associated ocean warming and acidification have profoundly transformed the marine realm (Pandolfi et al., 2011). Higher-than-usual seawater temperatures can cause coral bleaching, the breakdown of the symbiotic relationship between the coral host and its dinoflagellate endosymbionts (Symbiodinium spp.), and associated coral mortality (Hoegh-Guldberg, 1999). Ocean acidification is reducing carbonate ion availability in seawater and can depress calcification rates of calcifying organisms like corals (Langdon et al., 2000; Doney et al., 2009; Chan and Connolly, 2013). These global changes, coupled with local stressors such as pollution, overfishing, and outbreaks of crown-of-thorns starfish, have drastically altered coral cover and community composition at a global scale. In the last three decades, multiple mass bleaching events have decimated coral reefs worldwide including in 1998, 2010, and 2014–2017 (Eakin et al., 2016; Heron et al., 2016; Hughes et al., 2017, 2018). The Great Barrier Reef (GBR) is no exception with 50–80% coral mortality recorded on many northern reefs following the 2016 mass bleaching event (Great Barrier Reef Marine Park Authority, 2017), followed by another high mortality mass bleaching event in 2017. Climate models predict a <5% chance of reaching the Paris agreement target of limiting the global temperature rise to <2°C compared to pre-industrial times by 2100 (Raftery et al., 2017), and most coral reefs are forecasted to experience annual severe bleaching before the end of the century (van Hooidonk et al., 2016). Several observations of an increase in tolerance of coral bleaching after successive bleaching events suggest that adaptation and/or acclimatization are possible under certain conditions (Maynard et al., 2008; Berkelmans, 2009; Guest et al., 2012; Penin et al., 2013). Nevertheless, over 50% of the world's coral reefs has been lost in the last three decades, with the Caribbean having lost over 80% of its coral cover (50 Reefs, 2017)1, indicating that the rates of natural adaptation and acclimatization are overall insufficient to keep pace with the rate of environmental changes (van Oppen et al., 2017).
Active reef restoration is one way to assist the recovery of coral reefs that are degraded, damaged or destroyed. Reef restoration is still in its infancy and all of the few successful efforts so far occurred on a small spatial scale (e.g., Nakamura et al., 2011; Omori, 2011; Villanueva et al., 2012; Guest et al., 2014; dela Cruz and Harrison, 2017). Traditionally, locally sourced biological material is used for restoration based on the assumption that these populations are locally adapted and therefore most likely to survive (Breed et al., 2013). However, anthropogenic disturbances are rapidly changing the environment and shifting selection pressures (Becker et al., 2013), and locally sourced stock is therefore potentially mismatched with the altered environment. An effective restoration strategy should thus incorporate an understanding of present day ecological characteristics of species, characteristics of future available habitats, and adaptive potential of species (Becker et al., 2013). The use of non-local and climate resilient materials is controversial, but is gaining traction in wildland restoration (Jones and Monaco, 2009), revegetation (Sgrò et al., 2011; Breed et al., 2013), and coral reef restoration (Rau et al., 2012; van Oppen et al., 2017).
One possible way to improve the adaptive potential of species is via hybridization, which can increase genetic variation, break genetic correlations that constrain evolvability of parental lineages, and assist species to acquire adaptive traits (Hoffmann and Sgrò, 2011; Becker et al., 2013; Carlson et al., 2014; van Oppen et al., 2015; Hamilton and Miller, 2016; Meier et al., 2017). Hybridization can be conducted either via targeted crossing of individuals or species carrying desired phenotypic traits (e.g., high thermal tolerance) or via crossing between species with the goal of increasing genetic diversity and new variation for natural selection to act upon, and potentially generating hybrid vigor. The relative fitness of F1 hybrids (Figure 1) depends on whether there are additive (i.e., hybrids are of intermediate fitness between the parental species), dominant (i.e., hybrids are of equal fitness to the dominant parent species), over-dominant (i.e., hybrids are more fit than both parental species), under-dominant (i.e., hybrids are less fit than both parental species) gene effects, and/or maternal effects (i.e., hybrids are of equal fitness to their maternal parent species) (for review, see Lippman and Zamir, 2007; Li et al., 2008; Chen, 2013). Reciprocal hybrids are predicted to have equal fitness, except under maternal inheritance. With maternal effects, the fitness of the hybrids is directly affected by the fitness of the maternal parental species, regardless of the offspring's own genotype (Roach and Wulff, 1987; Bernardo, 1996). In the context of restoration, hybrid vigor which can be driven by dominant or over-dominant mechanisms, is a desirable outcome. The value of hybridization in enhancing fitness has been demonstrated in multiple cases. For instance, hybridization has provided genetic variance in morphology for adapting to changing environments in Darwin's finches (Grant and Grant, 2010), altered chemical defense of hybrid Brassicaceae plants and aided their survival through the Last Glacial Maximum (Becker et al., 2013), and facilitated extensive adaptive radiation in haplochromine cichlid fishes (Meier et al., 2017).
Figure 1. Possible relative fitness of reciprocal F1 hybrids (F1ab and F1ba) based on fitness of the parental species (Pa and Pb) and the driving mechanism. In the top graphs, parental species are assumed to differ in phenotype while the bottom graphs indicate a situation where parental species are similar. For further explanation, see text.
Hybridization is known to occur naturally in some scleractinian corals and has played an important role in the evolution and diversification of the genus Acropora (van Oppen et al., 2001; Willis et al., 2006). In the Caribbean, recent environmental degradation and massive population decline in Acropora cervicornis and Acropora palmata have favored hybridization and expansion of their F1 hybrid, Acropora prolifera (Fogarty, 2012). These hybrids either have equivalent or higher fitness relative to the parent species in most life history stages examined (Fogarty, 2012). In recent years, A. prolifera has been reported in increasingly high abundance in many reef locations (Fogarty, 2012; Japaud et al., 2014; Aguilar-Perera and Hernández-Landa, 2017) and the hybrid has expanded to marginal environments where parent species are absent (Fogarty, 2012).
Although interspecific hybridization is a potential tool to enhance restoration outcomes, it is often dismissed in restoration initiatives. Concerns raised include the possibility of outbreeding depression in later generations (i.e., F2, F3, backcross), and the loss of diversity through losing part of the parental species' genome (for review, see Hamilton and Miller, 2016). Most examples of outbreeding depression, however, are associated with the admixture of populations or species that are geographically distant, or when life history or phenological differences are large (Hwang et al., 2012; Whiteley et al., 2015). Outbreeding depression can also be transient and can be overcome by natural selection (Jones and Monaco, 2009; Aitken and Whitlock, 2013; Hamilton and Miller, 2016). Instead of reducing genetic diversity, hybridization may conserve diversity by protecting the parental genome from the risk of extinction, and can also increase genetic diversity by combining two divergent genomes within a single organism (Garnett et al., 2011). For example, hybridization has successfully enhanced genetic diversity, improved the population size and rescued the highly inbred, remnant population of Florida panther (Johnson et al., 2010) and the Mt. Buller mountain pygmy-possum (Weeks et al., 2017) from extinction (i.e., genetic rescue).
Here we investigate interspecific hybridization as a novel tool to increase genetic diversity and develop coral stock with increased climate resilience. Parental species were not chosen for their relative climate resilience, but based on our expert knowledge of the probability that they would cross-fertilize as well as their evolutionary relatedness. We examined the performance of hybrids from reciprocal crosses of two Acropora species pairs raised under ambient and elevated seawater temperature and pCO2 conditions, and assessed (1) whether prezygotic barriers exist in interspecific hybrids of Acropora corals from the GBR, and (2) whether hybrids show enhanced fitness and resilience compared to the purebreds. Four phenotypic traits (i.e., survival, recruit size, Symbiodinium uptake, and photochemical efficiency) were measured in hybrid and purebred offspring as proxies for fitness. Surviving hybrids and purebreds at the end of the experiment were transplanted to long-term grow-out tank for rearing with the aim to allow future assessment of their reproductive and backcrossing potential when they reach sexual maturity at ~4 years of age. We continued to monitor these survivors for survival and size during the grow-out period.
Materials and Methods
Coral Spawning, in Vitro Fertilization, and Experimental Design
A detailed timeline of the experiment, sampling and measurement of each trait is shown in Figure S1. Parental colonies were collected from Trunk Reef, central GBR, prior to full moon on 22nd Nov 2015 and maintained in flow-through aquaria of the National Sea Simulator (SeaSim) at the Australian Institute of Marine Science (AIMS). When signs of imminent spawning were observed (“setting,” i.e., where the sperm-egg bundles begin to protrude through the mouth of the polyps), colonies were isolated in individual tanks to avoid uncontrolled mixing of gametes prior to in vitro crossing. The five most profusely spawning colonies of each parental species were used for crossing to form (1) an Acropora tenuis × Acropora loripes cross, and (2) an Acropora sarmentosa × Acropora florida cross (Figure 2). These two species pairs were chosen to represent a phylogenetically divergent cross and a phylogenetically closely related cross. The phylogeny of Acropora spp. is divided into two distinct groups: the “early spawners” and the “late spawners,” where the “late spawners” spawn about 1.5–3 h before the other group (Fukami et al., 2000; van Oppen et al., 2001; Márquez et al., 2002). A. tenuis (early spawner) and A. loripes (late spawner) are phylogenetically divergent, while A. sarmentosa and A. florida (both are “late spawners”) are closely related and fall within the same phylogenetic clade (Fukami et al., 2000; van Oppen et al., 2001; Márquez et al., 2002). Little information is available from the literature about the relative resilience of these four parental species, but this has limited relevance for this study as our purpose was to increase genetic diversity (and thus adaptive potential) via hybridization, and not to conduct targeted breeding with species of known relative bleaching tolerance. Only two A. florida colonies spawned on the same night as A. sarmentosa, therefore, only two colonies were used for this species.
Figure 2. Experimental set up showing the two interspecific crosses [i.e., A. tenuis (T) × A. loripes (L) and A. sarmentosa (S) × A. florida (F)], the four resultant offspring groups from each cross (TT, TL, LT, LL; and SS, SF, FS, FF, respectively), larval settlement, and comparison of hybrid and purebred fitness under ambient and elevated conditions. The abbreviation of the offspring groups throughout this paper is that the first letter represents the origin of the eggs and the second letter the origin of sperm [e.g., SF is a hybrid of A. sarmentosa (S) eggs with A. florida (F) sperm]. The different colors used for the offspring groups in the figure reflect differently colored settlement plugs used for each offspring group.
Egg-sperm bundles of individual colonies were collected and eggs and sperm were separated using a 100 μm filter. Eggs were washed three times with filtered seawater to remove any residual sperm and placed in a 3 L bowl until crosses were set up (within 3 h). Sperm concentration of every colony was measured with a hemocytometer on a compound microscope with 40x magnification. Similar quantities of sperm from each conspecific colony were pooled to create a mixed sperm solution. For the hybrid crosses, the mixed sperm solution of the other parental species was added to the eggs of each interspecific colony. This method prevented intraspecific fertilization by possible remaining sperm that was not washed off the eggs (note that no self-fertilization was observed in any of the crosses performed). Fertilization was conducted under ambient conditions at a sperm concentration of 106 sperm mL−1. Three samples of 100 eggs were collected for each species as a self-fertilization test and a “no sperm” control. Each species pair cross produced four offspring groups, two purebreds and two hybrids (Figure 2). Embryos of each offspring group were then placed in rearing tanks for development under ambient conditions.
Fertilization Rates and Embryonic Development
Fertilization rates were assessed at 3.5 h, and embryonic development at 9, 15, 21, 33, 45, 57, and 93 h after sperm was added to the eggs. All embryos had reached planula stage and were ready to settle by 93 h. Triplicate samples of 100 embryos of each offspring group were collected and fixed in 4% formaldehyde. Developmental stages were assessed and counted under a dissecting microscope based on the stages described in Randall and Szmant (2009).
Larval Settlement and Symbiodinium Uptake
Prior to coral spawning, ceramic plugs of eight different colors were preconditioned in the outdoor SeaSim flow-through aquaria under ambient conditions for 6 weeks to develop crustose coralline algae (CCA) and a microbial biofilm to provide a larval settlement cue. Five days after fertilization, planula larvae of the eight offspring groups were each settled onto one assigned color of the pre-conditioned plugs under ambient conditions. Plugs of eight different colors were used so that each offspring groups could easily be identified and randomized in the experimental PVC trays holding the plugs (Figure 2). During settlement, Symbiodinium (i.e., algal symbionts) isolated from the parent colonies were added to achieve a final density of 2 × 106 cells mL−1 in each settlement tank. Larvae of each offspring group only received Symbiodinium from their parental species. To isolate the Symbiodinium, an ~6 cm fragment with three branches was removed from each parental colony with a bone cutter. Soft tissues of the fragment were then removed using an airbrush. The mixed soft tissues/seawater solution was collected and centrifuged at 200 g for 5 min to pellet the Symbiodinium. The Symbiodinium cells were resuspended and washed three times with filtered seawater before being added to the larvae. Symbiodinium uptake was assessed under a dissecting microscope prior to exposure to elevated conditions. Recruits (n = 20 per offspring group) were categorized as either with or without Symbiodinium.
Settled recruits were randomized and evenly distributed on 24 tailor-made PVC trays to rear under (1) ambient conditions of 27°C, 415 ppm pCO2, or (2) elevated conditions of ambient +1°C, 685 ppm pCO2 (Figure 2). Recruits for the elevated conditions were ramped to the target temperature and pCO2 from ambient at a rate of +0.2°C and + ~50 ppm per day. There were 12 replicate tanks for each of the two treatment conditions and tank positions in the experiment room were randomized (Figure 2). Every tank held one PVC tray with 20 plugs of each offspring group, with the exception of A. florida purebred (FF) and hybrid (FS) which had only 10 plugs due to fewer larvae being available. To avoid sediments from accumulating on top of the recruits, the trays were placed at an approximately 45° angle. Experimental conditions followed Davies Reef (18.83°S, 147.63°E) diurnal and annual temperature variations, a reef in proximity to Trunk Reef were the adult corals used for spawning were collected. A mixed marine microalgae diet of Isochrysis, Pavlova, Tetraselmis, Chaetocerous calcitrans, Thalassiosira weissflogii, and Thalassiosira pseudonana was fed to the recruits twice a day at a final concentration of ~5,000 cells mL−1 in the tank.
Survival and Recruit Size
Recruits from each tank were imaged using a high-resolution camera (Nikon D810) mounted on a quadpod with a waterproof case. Imaging was conducted fortnightly in the first 8 weeks of the experiment, thereafter every 4 weeks until 28 weeks. The numbers of surviving recruits were visually counted and recorded. Detailed images were taken at 28 weeks for size measurement. Recruit size was estimated as surface area of a circle from the measured recruit diameter since recruits were circular in shape and were not yet forming upright branches. Measurements were made using the software ImageJ and calibrated on the scale chart presented on every image. Recruits were maintained under the treatment conditions for 28 weeks. Surviving juveniles were thereafter relocated to long-term grow-out tanks to accommodate their larger size and maintained under ambient raw water (i.e., unfiltered seawater) to cater for higher feeding demand. Due to the small size of some recruits at 28 weeks and therefore difficulty to make comparisons, size was again measured at 1 year of age (i.e., about 5 months after all surviving recruits were moved to ambient raw water conditions) using the same measurement method. Furthermore, a set of photos of the median sized juvenile were taken at 2 years of age.
Photochemical efficiency (i.e., dark adapted maximum photosystem II quantum yield, Fv/Fm) was measured at week 28 as a proxy for coral health. Measurements were made using Imaging- Pulse Amplitude Modulation (I-PAM) derived by the software ImagingWin (v2.40b). Recruits (n = 15 per offspring group per treatment) were dark adapted overnight, and remained submerged in the treatment seawater during imaging. A recruit would only be measured if: (1) it was not obscured by filamentous algae, and (2) its size was no smaller than the software's area of interest requirement.
Automated controls of seawater chemistry were provided by SeaSim via the SCADA (Supervisory Control and Data Acquisition) system. Experimental conditions are summarized in Table 1. Seawater temperature and pH were recorded every hour using resistance temperature detector (RTD) and a pH probe (Tophit CPS471D). pCO2 was measured bi-weekly using a CO2 equilibrator calibrated to a standard gas of 500 ppm. Total alkalinity (AT) was measured using VINDTA calibrated to Dickson's Certified Reference Material. Salinity was measured weekly with an HACH IntelliCAL™ CDC401 Standard Conductivity Probe calibrated with IAPSO Standard Seawater. Seawater carbonate chemistry parameters, including Ωarag, DIC (dissolved inorganic carbon), , and were calculated using the measured values of seawater AT, pCO2, temperature and salinity, with the program CO2SYS (Lewis and Wallace, 1998 as implemented in Microsoft Excel by Pierrot et al., 2006).
Generalized linear mixed models (GLMM) (McCulloch and Neuhaus, 2013) for binomial data with logistic link functions were used to estimate the effects of treatment and offspring group on recruit survival at week 28. Analyses were conducted separately for the offspring groups of the A. tenuis × A. loripes cross and the offspring groups of the A. sarmentosa × A. florida cross using R Core Team (2016) with packages lme4 (Bates et al., 2014) and multcomp (Hothorn et al., 2008). In order to account for tank differences in the experimental design, a random tank effect was included in the models. Models were checked for overdispersion using a Chi-square test (Bolker et al., 2009) and goodness of fit using Akaike Information Criteria (Akaike, 1974). AIC of the GLMM of the A. tenuis × A. loripes cross was 397, the A. sarmentosa × A. florida cross was 331. Tukey's pairwise comparisons were then conducted and p-values were corrected using the Benjamini–Hochberg method (Benjamini and Hochberg, 1995). To obtain a visual overview of survival over time, longitudinal generalized linear models (GLM) for binomial data were used to estimate the survival for the offspring groups across all time points, and the combined hybrid offspring vs. purebred offspring. A summary table of the mean survival is provided in Table 7. Survival data were also analyzed with Cox proportional hazards regression as a comparison to GLMM. Results of the Cox regression were very similar to those of the GLMM and are shown in the Supplementary Methods and Results section (Table S1).
Statistical analyses of size were conducted separately for the offspring groups of the A. tenuis × A. loripes cross and the offspring groups of the A. sarmentosa × A. florida cross at 28 weeks and at 1 year of age. For the 28-week time point, the absence of growth in a large number of samples under elevated conditions resulted in non-normality of the data, and non-parametric Kruskal–Wallis tests were undertaken followed by Dunn's pairwise comparisons (Dunn, 1964). P-values for the multiple pairwise comparisons were adjusted with the Benjamini–Hochberg method. For the 1 year time point, due to the absence of survivors in some offspring groups, not all size comparisons could be undertaken. Offspring groups with no or less than three survivors were excluded from the analyses. The remaining size data were normally distributed (tested using Shapiro–Wilk tests; Shapiro and Wilk, 1965) and variances were homogeneous (tested by Levene's tests; Levene, 1960) and five pairwise comparisons were undertaken using combined variance t-tests. For the A. tenuis × A. loripes cross, three t-tests were possible for offspring groups that were previously exposed to ambient conditions (note they have been relocated to long-term grow-out tank under raw ambient seawater after 28 weeks). The p-values of these comparisons were adjusted using the Benjamini–Hochberg method. The above analyses were run in R (version 3.3.1). A summary of the mean sizes of the recruits is shown is Table 7.
Symbiodinium Uptake and Photochemical Efficiency
Generalized linear models (GLM) (McCulloch and Neuhaus, 2013) were used to test the effect of offspring group on rates of Symbiodinium uptake, which was treated as a binomial distributed variable (i.e., Symbiodinium taken up/not taken up). Treatment was not included in this model as Symbiodinium uptake was assessed prior to the start of treatment. For photochemical efficiency (i.e., dark adapted yield, Fv/Fm), generalized linear models were also used to test the effects of offspring group and treatment on the response. Tukey pairwise comparisons were then used and the p-values were adjusted with the Benjamini–Hochberg method. These analyses were run with R packages lme4 (Bates et al., 2014) and multcomp (Hothorn et al., 2008).
Spawning Time, Fertilization Rates, and Embryonic Development
The date and time of spawning of the Acropora spp. used in this study are summarized in Table 2. The A. tenuis × A. loripes cross was conducted on the 6th day after the full moon, where A. tenuis spawned at ~19:00–19:30 and A. loripes at ~21:45. The A. sarmentosa × A. florida cross was conducted on the 7th day after the full moon, where A. sarmentosa spawned at ~20:30–20:45 and A. florida at ~21:15–21:30. Fertilization rates of all but one hybrid offspring group were high (averaged 93%) (Figure 3). Hybrid LT had lower fertilization rates (averaged 79%) compared to all other offspring groups. No fertilization was observed in the “no-sperm” control and self-fertilization tests. Purebred and hybrid embryos developed normally and reached the planula stage 93 h after fertilization (Figure S2). The hybrid LT also had a slower initial embryonic development rate with the majority of the LT embryos being at the 2–4 cell stage at 3.5 h after fertilization, while embryos of all other offspring groups were at the 8–16 cell stage (Figure S2). From 9 h onwards, however, all offspring groups developed at similar rates.
Figure 3. Fertilization rates of the offspring groups from (A) the Acropora tenuis (T) × Acropora loripes (L) cross, and (B) the Acropora sarmentosa (S) × Acropora florida (F) cross. The abbreviation of the offspring groups is that the first letter represents the origin of the eggs and the second letter the origin of sperm, where TL, LT, SF, FS are hybrids and TT, LL, SS, FF are purebreds.Values are mean and error bars represent 95% CI calculated using the angular transformed data back-transformed into percentages.
Overall, maternal effects were observed in the hybrid offspring groups of the A. tenuis × A. loripes cross and over-dominance in the A. sarmentosa × A. florida cross, with some variations between treatment conditions (Figure 4). Offspring groups differed significantly for survival both in the A. tenuis × A. loripes cross (GLMM, χ2 = 252.2, df = 3, p < 0.001), and the A. sarmentosa × A. florida cross (GLMM, χ2 = 32.2, df = 3, p < 0.001). The values present below are mean survival and the associated 95% confidence intervals are shown in Table 3. For the A. tenuis × A. loripes cross, survival of hybrid LT (49%) and purebred LL (46%) was higher than that of TT (13%) and TL (16%) under ambient conditions (p < 0.001 for all) (Tables 3, 4, 7). Under elevated conditions, survival of hybrid LT (41%) and purebred LL (36%) was also higher than that of TT (7%) and TL (23%) (Tables 3, 4, 7). Survival of hybrids was similar to that of their maternal parental purebred offspring. Under elevated conditions, survival of hybrid TL (23%) was also higher than that of purebred TT (7%) (p < 0.001; Tables 3, 4, 7).
Figure 4. Survival of the offspring groups of (A) the Acropora tenuis (T) × Acropora loripes (L) cross, and (B) the Acropora sarmentosa (S) × Acropora florida (F) cross under ambient and elevated conditions across 28 weeks. The abbreviation of the offspring groups is that the first letter represents the origin of the eggs and the second letter the origin of sperm. Lines represent the estimates of the longitudinal generalized linear models.
Table 3. Mean survival, SE, as well as lower and upper 95% CI of offspring groups from the Acropora tenuis (T) × Acropora loripes (L) cross and the Acropora sarmentosa (S) × Acropora florida (F) cross under ambient and elevated conditions.
Table 4. Tukey's pairwise comparisons of survival between the offspring groups from the Acropora tenuis (T) × Acropora loripes (L) cross and the Acropora sarmentosa (S) × Acropora florida (F) cross following generalized linear mixed models.
For the A. sarmentosa × A. florida cross, survival of both hybrid SF (51%) and FS (53%) was higher than that of the purebred FF (31%) (p = 0.014, p = 0.006, respectively) and SS (35%) (p = 0.022, p = 0.007, respectively) under ambient conditions (Tables 3, 4, 7). Under elevated conditions, only hybrid FS (32%) had higher survival than purebred SS (18%) (p = 0.007; Tables 3, 4, 7). When combining the data for all hybrid and purebred offspring groups for an overall comparison, hybrid offspring had a consistently higher survival than purebred offspring (Figure S3).
Treatment had a significant effect on survival of both the A. tenuis × A. loripes cross (GLMM, χ2 = 26.9, df = 1, p < 0.001), and the A. sarmentosa × A. florida cross (GLMM, χ2 = 13.6, df = 1, p < 0.001). Tukey pairwise comparisons suggest that TT, SS, SF, and FS had lower survival under elevated conditions compared to ambient conditions at week 28 (p = 0.025, 0.002, 0.007, 0.015, respectively; Table 5).
Table 5. Tukey's pairwise comparisons of treatment effect within an offspring group from the Acropora tenuis (T) × Acropora loripes (L) cross and the Acropora sarmentosa (S) × Acropora florida (F) cross following generalized linear mixed models.
For the A. tenuis × A. loripes cross, treatment had a significant effect on recruit size (Kruskal–Wallis, χ2 = 33.6, df = 1, p < 0.001) but offspring group did not (Kruskal–Wallis, χ2 = 6.9, df = 3, p = 0.096; Figure 5). For the A. sarmentosa × A. florida cross, treatment also had a significant effect on recruit size (Kruskal–Wallis, χ2 = 38.2, df = 1, p < 0.001). Offspring group had a significant effect on size under ambient conditions (Kruskal–Wallis, χ2 = 18.2, df = 3, p < 0.001), but not under elevated conditions (Kruskal–Wallis, χ2 = 1.0, df = 3, p = 0.793). Under ambient conditions, the mean size of hybrids FS (41 mm2) and SF (43 mm2) was larger than that of the purebred SS (16 mm2) (z = 3.19, p = 0.003; z = 3.56, p = 0.001, respectively), but not different in size from FF (56 mm2; Table 7).
Figure 5. Boxplots showing the size of the Acropora offspring groups at 28 weeks since treatment began from (A) the Acropora tenuis (T) × Acropora loripes (L) cross and (B) the Acropora sarmentosa (S) × Acropora florida (F) cross. The first letter of the offspring groups' abbreviation represents the origin of the eggs and the second letter the origin of sperm. The horizontal bars represent median values, box length represents the interquartile range, and the small circles denote unusual points.
At the 1-year time point, several offspring groups no longer had survivors (Figure 6, Table 7). Note that the treatment condition in this section refers to the treatment conditions that the recruits were exposed to during the 28 week period following settlement, but that they were transferred to long-term grow-out tanks with ambient raw (i.e., unfiltered) seawater afterward. For recruits that were previously under ambient conditions, there were no survivors of purebreds TT and FF, while all hybrid groups had survivors. The mean size of LT (362 mm2) and LL (366 mm2) offspring was larger than that of TL offspring (47 mm2) (t-test, p = 0.008, 0.015, respectively, Tables 6, 7). The size of the LT hybrids was the same as that of the maternal parent species LL (i.e., maternal effect). The mean size of FS hybrids (304 mm2) was larger than that of the pure breds SS (30 mm2) (t-test, p = 0.004, Tables 6, 7). The mean size of hybrid SF (245 mm2, average of 2 recruits) was also relatively larger than SS (30 mm2), however, statistical comparison was not possible due to the low sample size for SF (n = 2; Table 7). For recruits that were previously under elevated conditions, there were no survivors of purebreds TT and SS as well as hybrids TL and SF. The mean size of the hybrid LT recruits (326 mm2) was the same as that of the maternal parent species LL (290 mm2) (Tables 6, 7). Median sized survivors of hybrid and purebred juveniles at 2 years of age and are shown in Figure S4.
Figure 6. Boxplots showing the size of the Acropora offspring groups at 1 year of age (i.e., ~5 month since relocation to long-term grow-out tank under raw ambient seawater. (A) The Acropora tenuis (T) × Acropora loripes (L) cross and (B) the Acropora sarmentosa (S) × Acropora florida (F) cross. The first letter of the offspring groups' abbreviation represents the origin of the eggs and the second letter the origin of sperm. Where no data are presented there were no survivors in that offspring group. The horizontal bars indicate the medians, box length indicates the interquartile range, and the small circles indicate unusual points. Images below the graphs show examples of median size recruits of the offspring groups reared under ambient conditions in the experiment, and the number of survivors of each offspring group.
Table 6. Results of t-tests comparing size at the 1-year time point for remaining offspring groups of the Acropora tenuis (T) × Acropora loripes (L) cross, and the Acropora sarmentosa (S) × Acropora florida (F) cross.
Symbiodinium Uptake and Photochemical Efficiency
There was no significant difference in Symbiodinium uptake between the offspring groups of the A. tenuis × A. loripes cross (GLM, χ2 = 3.25, df = 3, p = 0.354) or the offspring groups of the A. sarmentosa × A. florida cross (GLM, χ2 = 5.35, df = 3, p = 0.148; Figure S5, Table 7). For the A. sarmentosa × A. florida cross, neither treatment nor offspring groups had a significant effect on photochemical efficiency (Treatment: GLM, χ2 = 0.51, df = 1, p = 0.477; offspring group: GLM, χ2 = 4.28, df = 3, p = 0.233). For the A. tenuis × A. loripes cross, treatment had a significant effect on photochemical efficiency (GLM, χ2 = 6.87, df = 1, p = 0.009) but offspring groups did not (GLM, χ2 = 2.43, df = 3, p = 0.488; Figure S6). Tukey pairwise comparisons show that purebreds TT and LL under elevated conditions had lower photochemical efficiency than their counterparts under ambient conditions (p = 0.035, 0.002, respectively).
The results of the various traits measured are summarized in Table 7.
Limited Prezygotic Barriers to Interspecific Hybridization in Acropora Corals
To understand the value of hybridization for coral reef restoration, it is important to establish whether prezygotic barriers exist. Interspecific hybridization among Acropora spp. has been shown to occur in experimental crosses, with varying degrees of prezygotic barriers (Willis et al., 1997; Van Oppen et al., 2002; Fogarty et al., 2012; Isomura et al., 2013). Among multiple pairs of Acropora spp. from the central GBR, crossing resulted in eight pairs with high fertilization (50–80%), seven pairs with moderate fertilization (10–50%) and three pairs with low fertilization (3–10%) (Willis et al., 1997; Van Oppen et al., 2002). The high fertilization rates and normal embryonic development of the interspecific hybrids produced in this study indicate prezygotic barriers are limited in these species pairs. This was unexpected in the case of the A. tenuis × A. loripes cross which involved an “early spawner” and a “late spawner.” These “early spawners” and “late spawners” are believed to have diverged 6.6 Mya (Fukami et al., 2000). We hypothesize that our observations can be explained by the fact that the gametes of these species do not normally encounter one another in the field due to a 2 h difference in spawning times, and selection on prezygotic barriers has therefore been absent. Conversely, A. sarmentosa and A. florida are phylogenetically closely related, occur sympatrically and spawned ~30 min to 1 h apart. Our results indicate a prezygotic barrier has not evolved to maintain reproductive isolation of these two species either, and that hybridization may occasionally occur in nature.
Lower fertilization in one direction in Acropora hybrid crosses, as was observed for hybrid LT, is not uncommon (Fogarty et al., 2012; Isomura et al., 2013). The likelihood of A. palmata eggs being fertilized by A. cervicornis sperm, for instance, is smaller than the likelihood of A. cervicornis eggs being fertilized by A. palmata sperm (Fogarty et al., 2012). The lower fertilization rate of the hybrid LT, however, did not affect recruit size or survival of this offspring group. The slight delay in embryonic development as observed in the hybrid LT was similar to observations for another Acropora cross in Japan (Isomura et al., 2013). This delay, however, was only limited to one early time point and no aberrant development was observed in the hybrids at any time point. The results of this and previous studies suggest that the degree of prezygotic barriers varies between Acropora species, and a range of species with limited prezygotic barriers can be used for hybridization with the aim to enhance climate resilience. Interspecific hybridization may also be applied to several other coral genera. Experimental crosses have successfully hybridized species within the genera Montipora and Platygyra (Willis et al., 1997), but were unsuccessful for species in the genus Ctenactis (Baird et al., 2013). Future studies to test the success of interspecific hybridization in additional coral genera will be valuable to determine the extent to which this approach for coral reef restoration can be applied.
Positive Effects of Hybridization Were Observed in Some F1 Hybrids
Given only limited prezygotic barriers exist, we explored whether hybrid offspring had increased resilience and may be used to enhance coral reef restoration efforts. If hybrids are comparatively resilient, interspecific hybridization may be combined with methods being developed for deploying coral larvae or recruits onto reefs requiring restoration (e.g., Nakamura et al., 2011; Omori, 2011; Villanueva et al., 2012; Guest et al., 2014; dela Cruz and Harrison, 2017). Overall, maternal effects were observed in hybrids of the A. tenuis × A. loripes cross and over-dominance in hybrids of the A. sarmentosa × A. florida cross, with some variations between traits and treatment conditions. Possible benefits of hybridization in enhancing reef restoration can be observed in both crosses (Table 7). In the A. tenuis × A. loripes cross, hybrids of both directions exhibited ~16–34% higher survival than purebred A. tenuis under conditions with elevated temperature and pCO2 (Table 7), suggesting hybrids have higher climate resilience than the purebred species. Both purebred species also showed reduced photochemical efficiency under elevated compared to ambient conditions while both hybrid species did not. Furthermore, purebred A. tenuis had no survivors at the 1-year time point, yet hybrids from both directions survived. For the A. sarmentosa × A. florida cross, hybrids of one or both directions showed ~14–22% higher survival and were larger in size than both or one parental purebred species (Table 7). One hybrid offspring group (FS) had survivors in both ambient and elevated conditions at the age of 1 year, while both purebred species had no survivors in one of these conditions (Table 7). The FS hybrid was also 10 times larger in size than the only surviving purebred species (A. sarmentosa) at 1 year of age under ambient conditions (Table 7).
Across all traits measured, hybrids were either equivalent to or more fit than at least one parent, and none of the hybrids performed worse than both parents. These patterns are similar to those seen in some other comparisons. The natural hybrid A. prolifera in the Caribbean (Fogarty, 2012) and experimentally produced hybrids of A. millepora × A. pulchra (Willis et al., 2006) had equivalent or higher fitness compared to their parental species. Experimentally produced hybrids of A. millepora × A. pulchra grow larger in size than purebreds in the reef-flat environment (Willis et al., 2006). In this study, photochemical efficiency was the same between offspring groups in the same treatment, suggesting that (1) the observed differences in recruit size of the offspring groups were unlikely caused by carbon translocation from Symbiodinium, and (2) there was no coral host effect on photochemical efficiency of the Symbiodinium.
Hybrid Fitness and Its Relevance to Coral Reef Restoration
A comprehensive assessment of the value of interspecific hybridization to coral reef restoration requires multi-generation fitness examinations of hybrids and backcrosses. Such an assessment will require years given the long generations time of corals (3–7 years to reach reproductive maturity). The present study is one of the few studies that examines the long-term fitness of F1 hybrids and provides detailed assessments from fertilization to embryonic development, Symbiodinium uptake, photosynthesis efficiency, survival and size. The results provide evidence that hybridization may have value to reef restoration. From the restoration point of view, hybridization increases genetic variation which can potentially enhance adaptive capacity and release a population from adaptive limits (Hoffmann and Sgrò, 2011; Becker et al., 2013; Carlson et al., 2014; van Oppen et al., 2015; Hamilton and Miller, 2016; Meier et al., 2017). In this study, genetic diversity would have increased in the F1 hybrids and positive effects on survival and recruits size were observed in some cases. Furthermore, none of the hybrid offspring groups performed worse than the purebreds across all traits, suggesting that there was no negative effect of hybridization in the F1. Higher survival and larger recruit size as observed in some hybrids can enhance reef restoration initiatives by reducing post-settlement mortality. Moreover, reproductive maturity of a coral is related to its size (Soong and Lang, 1992; Smith et al., 2005). Corals that achieve a large size earlier can begin to reproduce sooner, which may further assist the recovery of degraded reef systems.
In the A. tenuis × A. loripes cross, maternal effects were observed in fitness. Hybrid survival and size were similar to that of the maternal purebred species, although it exceeded purebred values in some occasions/conditions. Maternal effects have previously been shown for survival of interspecific hybrid larvae from an A. florida × A. intermedia cross (Isomura et al., 2013), and for thermal tolerance and gene expression levels of intraspecific A. millepora hybrid larvae from a higher and lower latitude cross (Dixon et al., 2015). It is unclear whether the observed fitness for the F1 hybrids from our study is due to nuclear or cytoplasmic maternal effects. If survival and size are governed by nuclear maternal effects, F2 hybrids of both directions will have similar fitness to each other, which will be different from their maternal parent. If survival and size are controlled by cytoplasmic maternal effects, fitness of the F2 hybrids will follow maternal fitness (Roach and Wulff, 1987; Bernardo, 1996). In this case, species that are known to carry desirable traits under climate change may therefore be ideal candidates as a source of eggs for creating coral stock for restoration via interspecific hybridization.
When selecting species pairs for hybridization to facilitate reef restoration, both targeted crossing with species or individuals that carry phenotypic traits of value under climate change (e.g., high thermal tolerance) and non-targeted crossing between species could be considered. Species with high climate resilience (e.g., A. loripes in the present study) may be useful for targeted hybridization efforts. Alternatively, non-targeted crossing among related species could be used to generate hybrid vigor and increase genetic diversity for future adaptation.
Knowledge Gaps and Future Studies
This study provides the first steps toward the assessment of interspecific hybridization as an approach to create coral stock with augmented climate resilience. While our findings are supportive of this novel strategy, additional research is required. Three important outstanding questions are: (1) The fitness of hybrids vs. purebreds in the field. While laboratory studies are ideal to investigate the responses of corals to one or two specific stressor(s), corals in the wild are subjected to other selection pressures difficult to simulate in the laboratory. An important next stage of this research would involve outplanting the hybrids and purebreds to the field and monitoring their relative fitness. (2) The reproductive and backcrossing potential of F1 hybrids. Isomura et al. (2016) have shown that experimentally produced A. intermedia × A. florida F1 hybrids were fertile and able to produce F2 offspring with high fertilization rates. Transgressive segregation can happen in F2 hybrids, where segregating variation of parental species recombines in hybrids at multiple loci to produce extreme phenotypes and may result in some F2 hybrids with extremely high fitness (for review, see Hamilton and Miller, 2016). Conversely, outbreeding depression may become apparent in the F2 generations and result in hybrids with low fitness. F1 hybrids of the A. intermedia × A. florida cross were able to backcross with either both parent species or the maternal parent species only, depending on the direction of the hybrid cross (Isomura et al., 2016). In the Caribbean, molecular evidence has shown unidirectional gene flow from A. palmata into A. cervicornis, suggesting that their hybrid A. prolifera is fertile and able to backcross with at least one parental species (Vollmer and Palumbi, 2002, 2007). Current knowledge on fertility of F1 coral hybrids remains limited and future studies in the area will be invaluable. (3) The fitness of advanced generation hybrids and backcrosses. If F1 hybrids are fertile and sexual reproduction is successful, high fitness will have to be maintained in the F2, backcrosses and advanced generation hybrids for interspecific hybridization to be beneficial to reef recovery and resilience in the long-term.
F1 hybrids can theoretically propagate via asexual reproduction, and via sexual reproduction with other hybrids or the parental species. The likelihood of the latter depends on the spawning time of the F1 hybrids and the parental species. The A. tenuis × A. loripes cross in the present study had 2 h difference in spawning time and the A. sarmentosa × A. florida cross had 30 min to 1 h difference. While the spawning time of the hybrids remains unknown until they reach reproductive maturity, Isomura et al. (2016) showed that F1 Acropora hybrid spawned at the same time as their maternal parental species. This suggests that the F1 in the present study will likely be able to at least backcross with the maternal parent species for the A. tenuis × A. loripes cross, and potentially with both parental species for the A. sarmentosa × A. florida cross due to closer spawning time. Asexual reproduction (i.e., fragmentation, polyp bail-out) is a common reproductive strategy of broadcast spawning scleractinian corals (Highsmith, 1982; Sammarco, 1982) and F1 hybrids may also persist in the wild via this method. A. prolifera, the natural F1 hybrid in the Caribbean for example, is known to persist and colonize large reef areas through asexual reproduction (Irwin et al., 2017).
In sum, it is likely that interspecific Acropora hybrids are able to propagate over extended periods of time, either sexually, asexually, or via both reproductive methods. Before hybrids can safely be used as stock for restoration, however, it must be demonstrated that the risk of this strategy is low by showing that the fitness of later generations remains equal or superior to that of the parental species in the wild.
WC, MvO, LP, and AH: designed the experiment; MvO: developed the concept for this study; WC and LP: conducted the experiment and collected the data; PM, WC, and AH: undertook statistical analyses; WC and MvO: wrote the manuscript and all authors contributed to the final edited version of the manuscript.
Conflict of Interest Statement
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
This study was funded by the Paul G. Allen Family Foundation and the Australian Institute of Marine Science (AIMS). We thank staffs of the National Sea Simulator at AIMS for technical support. Thanks are also extended to M. Nordborg for her contribution to analysing the coral embryos, K. Damjanovic, R. Alfred, and L. Blackall for help with coral spawning work, and M. Shanahan for seawater alkalinity measurements. WC acknowledges receipt of the University of Melbourne International Research Scholarship and Fee Remission Scholarship.
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmars.2018.00160/full#supplementary-material
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Keywords: hybridization, restoration, coral reefs, climate change, Acropora, assisted evolution, genetic rescue, hybrid vigor
Citation: Chan WY, Peplow LM, Menéndez P, Hoffmann AA and van Oppen MJH (2018) Interspecific Hybridization May Provide Novel Opportunities for Coral Reef Restoration. Front. Mar. Sci. 5:160. doi: 10.3389/fmars.2018.00160
Received: 19 January 2018; Accepted: 23 April 2018;
Published: 14 May 2018.
Edited by:Anastazia T. Banaszak, Universidad Nacional Autónoma de México, Mexico
Reviewed by:James R. Guest, Newcastle University, United Kingdom
Mathieu Pernice, University of Technology Sydney, Australia
Copyright © 2018 Chan, Peplow, Menéndez, Hoffmann and van Oppen. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Wing Yan Chan, email@example.com

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—Chamatkar Sandhu (@SandhuMMA) June 19, 2020
Tyson finished his career with a record of 50 wins (44 knockouts) against 6 losses after heavy, back-to-back knockout defeats in 2005.
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domain: medical


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Last Updated on August 6, 2015 – 11:30 AM CDT
The setting Panhandle sun lights up the memorial on the west side of the Moore County Courthouse on which the names of 19 firefighters who died 59 years ago are etched. Nineteen men with families, some of whom gathered Wednesday, as they do each year, to remember the sacrifices those men made July 29, 1956.
Dumas Fire Chief Paul Jenkins led the service to honor the men who died battling a fire at the Shamrock Oil Refinery, now Valero. The men were part of the Dumas and Sunray fire departments and the Shamrock Industrial Fire Brigade, and it is still one of the largest losses of firefighter lives in U.S. history. Forty other men were severely burned.
You can watch a video of the memorial below.

domain: history


Electric scooters have become the latest craze in urban transportation, and for good reason. Niu Scooter Electric E150. They are eco-friendly, cost-effective, and convenient. Niu Electric Scooters are a standout among the many brands on the market. They have a sleek design, great features, and high customer satisfaction. In this ultimate review, we’ll take a deep dive into everything you need to know about Niu Electric Scooters.
We have everything you need, from the most recent models to their unique features. We’ll also explore the pros and cons of these scooters, and what makes them stand out from the competition. You can read on to learn everything you need about Niu Electric Scooters, no matter if you are a veteran rider or just starting out.
If you have ever considered purchasing a Niu electric scooter, it is a great idea. Here’s why:
Electric scooter green from Niu – Niu Scooter Electric E150
Electric scooters can be a green and clean mode of transport. Electric scooters aren’t as polluting the environment as cars, and are better for the environment that driving.
They are also cheaper to run than cars. This means you can save money on gas and insurance.
Electric scooters are safer than cars because they’re smaller, lighter and more maneuverable. They don’t have the same blind spots that cars do, so you can see around corners and through intersections much better than in a car.
You can travel quickly and far.
An electric scooter can travel up to 25 miles per hour, which is much faster than you can pedal a bicycle. This means that you can go much farther in a shorter amount of time, without getting tired or sweaty.
Electric scooters are also easier to ride uphill than bicycles because they don’t require as much effort from the rider. If you’re heading up a steep hill and your legs get tired from pedaling, simply switch over to electric mode!
Electric scooters are also safer than bicycles. You are less likely to be hit by someone behind you if you’re on an electric scooter.
An electric Niu scooter can help you save money on gasoline.
Electric scooters are much cheaper than gasoline-powered ones. This means that you will save money over the long term. Electric scooters have lower fuel costs, maintenance costs and insurance premiums than their counterparts. These can add up to thousands over the life of your scooter’s ownership.
When it comes to performance, electric scooters are no slouch. Despite their compact size, they can reach impressive speeds, often up to 15-20 mph. Some models have powerful motors that can handle steep hills easily. Additionally, electric scooters are incredibly maneuverable, making them perfect for navigating through crowded streets and tight spaces.
But, the battery life is one of the most important aspects that an escooter’s performances. It is not worth having a scooter that can only go for a few seconds. Modern electric scooters are equipped with lithium-ion battery packs that provide long-lasting power. Depending on the model, you can expect anywhere from 10 to 30 miles of range on a single charge. Additionally, many escooters feature quick-charging technology, allowing you to juice up the battery in just a few hours.
Electric scooters are also more environmentally friendly than gas-powered ones. Electric scooters don’t emit exhaust so they don’t pollute the environment or contribute to greenhouse gas emission. It also doesn’t use fossil fuels, which means you won’t be contributing to climate change by using one! Niu Scooter Electric E150.
Electric scooters are fun!
It’s a great way for people to meet and have fun with electric scooters. They are easy to control and learn, making them great for children. However, adults love them too. They can be used for racing, which is a popular hobby among electric scooter enthusiasts.
Electric scooters are available in many different styles, from ones with only one wheel to ones that look like motorcycles. You can choose from aluminum or carbon fiber as they are made of a variety materials.
The electric motors of an electric scooter are powered by rechargeable batteries. They usually have a top speed of around 15 miles per hour, which is fast enough to get you where you need to go but slow enough that they’re safe for kids.
Although electric scooters can be very easy to use, they are still quite dangerous. You should have the proper training and protective gear if you intend to ride one. It’s also important to keep in mind that these scooters aren’t legal everywhere, so check your local laws before purchasing one.
Niu Electric scooter maintenance
Electric scooter maintenance involves several tasks, such as checking the brakes, tires, and battery. Regular inspection of the brakes is necessary to ensure that they work properly. Faulty brakes can cause accidents. Check the tires for wear and tear and inflate them to the recommended pressure. It is important to maintain the battery in order to ensure that the scooter is reliable and has a long range.
Other tasks involved in electric scooter maintenance include cleaning the scooter, lubricating the moving parts, and checking the lights. Regular cleaning prevents dirt and debris buildup, and lubricating moving parts will ensure they run smoothly. Lights are important for safety when riding at night, so they should be checked to ensure they are working correctly.
Regular maintenance is not enough. Electric scooters can also need occasional repairs. This could include the replacement of the battery, motor, and other parts that have become worn over time. To ensure that repairs are done properly, it is important to hire a qualified technician.
Owning a Niu electric scooter is a great way to get around town
An electric scooter can be a great way of getting around the city. They’re fun to ride, easy to maintain and store, and easy to park.
These scooters are ideal for students and commuters, but also for those who simply want to move around the city. They’re fun to ride, easy to maintain and store, and easy to park.
These are also great for people with disabilities.
Electric scooters are fun to ride, easy to maintain and store, and easy to park. These scooters are a great way to help people with disabilities get around their community and enjoy it.
Conclusion: Niu Scooter Electric E150
We hope you enjoyed reading about the many benefits of owning an electric bike. If you’re interested in getting one for yourself and want to learn more about what they have to offer, check out our guide on how to choose the best model for your needs!


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Boutique residential developer Regal London is aiming to double the amount of London property it sells in the GCC or Gulf Cooperation Council nations and has opened a Dubai office to help it achieve that goal.
In an interview with Abu Dhabi based website The National, Behrang Jalali, Director, Middle East and North Africa adds that the company’s GCC sales have remained steady at around 12% on average for the last five years, although the Brexit vote dampened interest for a time. However last year 18.5% of its sales were to investors from the GCC. Jalali added that the Dubai office has been opened to “court the growing pool of investors seeking cheap deals ahead of Brexit”.
Regal London has annual revenues around £225m and currently has five projects in London. Two schemes in King’s Cross and Queen’s Park are almost complete, two in West Kensington and St John’s Wood are due for completion in 1H 2019, while a project in Shoreditch is targeted for delivery in 2020.
Jalali told The National: “Those buying in dollar-denominated currencies stand to gain more from the weak pound and we expect this to continue.
“We think prices will stabilise and rise again in the years ahead. For investors, it’s good to diversify and London has bigger upside potential [than downside risks].”
Regal London’s Gulf sales drive is another small but significant vote of confidence in the long term prospects for the London property market. It also underlines the growing willingness of Asian and Middle Eastern investors to bet on Brexit.

domain: geography


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Washington, Cal meet in conference play
Washington (11-5, 1-2) vs. Cal (7-8, 1-1)
Haas Pavilion, Berkeley, California; Saturday, 8 p.m. EST
BOTTOM LINE: Pac-12 foes meet as Washington takes on Cal. Both teams last saw action on Thursday. Cal won at home over Washington State 73-66, while Washington came up short in a 61-55 game at Stanford.
LEADING THE WAY: Cal’s Matt Bradley has averaged 17.7 points and 5.1 rebounds while Grant Anticevich has put up 9.1 points and 5.7 rebounds. For the Huskies, Isaiah Stewart has averaged 18.5 points and 8.8 rebounds while Jaden McDaniels has put up 13.6 points and 6.1 rebounds.MIGHTY MATT: Bradley has connected on 36.1 percent of the 72 3-pointers he’s attempted and has gone 6 for 26 over his last five games. He’s also made 85.7 percent of his foul shots this season.
UNDEFEATED WHEN: Cal is a perfect 6-0 when the team makes six or more 3-pointers. The Golden Bears are 1-8 when the team hits fewer than six threes.
ASSIST RATIOS: The Huskies have recently used assists to create baskets more often than the Golden Bears. Cal has 21 assists on 64 field goals (32.8 percent) across its past three outings while Washington has assists on 26 of 66 field goals (39.4 percent) during its past three games.
STIFLING DEFENSE: Washington has held opposing teams to 37 percent shooting from the field this year, the 10th-lowest percentage among all Division I teams. Over their last five games, the Huskies have held opposing shooters to 32.4 percent.


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September 19, 2020
Promoting Positivity in the Workplace
Submitted by the USPS Employee Assistance Program
Do you ever have one of those days where everything seems to be going wrong and you just can’t catch a break? Sometimes, just watching the news or scrolling through social media can make a person feel defeated. Negativity is like a sprawling weed. It starts in one area and can spread quickly and easily to other areas of our lives, in our homes and in our workplaces.
There are many things in this world over which we do not have control, but we do have control over how we react to situations. Having a positive outlook is an inside job that starts with you, your thoughts and your actions. The good news is positivity can be contagious.
There are numerous studies that support how positive work environments encourage an increase in effectiveness, productivity and employee engagement. You can directly affect your workplace environment by choosing to be positive. Those in supervisory roles determine what is acceptable and not acceptable in the workplace by how they communicate and manage their personal interactions with others.
As a supervisor, you have the ability to encourage and promote positive behavior in the workplace. By engaging in positive and respectful behavior in your interactions with your colleagues, superiors and employees, you can model what healthy and positive workplace interaction should look like.
Greeting others by making eye contact and saying “good morning” or “good afternoon” goes a long way. Encourage an environment where people move forward. Acknowledge when mistakes are made, make the appropriate corrections, then move on.
In many instances, we spend more of our time at work than we do with our families. It is important to foster a positive work environment and model positive behavior and interactions with others. Focus on the aspects of your job that bring you fulfillment. Being passionate about your job and having empathy and compassion for your colleagues help improve workplace morale.
Another great way to promote positivity in the workplace is by showing gratitude and appreciation toward others. This encourages engagement. Being authentic and sincere in your interaction with others is a wonderful way to make others feel valued. Acknowledging others for a job well done or going above and beyond is important.
The saying that a person who is appreciated will always do more than what is expected is true. It is human nature for people to want to feel appreciated. Employees and colleagues who feel valued will want to engage as team players; this will increase productivity, attendance and cooperation.
Being friendly and approachable toward your employees and colleagues will help improve work relationships and create a more positive environment. Seek to find something likable in everyone around you. Differences in personalities, work ethics and cultural backgrounds can lead to a stressful work environment. Challenge yourself to find the good in every person with whom you come in contact.
Listening and being open to others’ opinions, ideas and solutions without judgment encourage employees to speak up and feel heard as valued members of the team. Model and encourage an environment where it is okay to agree to disagree in a respectful manner to help promote a positive work atmosphere. By changing the way you respond to disagreements, you can model positive and effective communication in the workplace. Below are a few examples:
As a supervisor, your attitude and work ethic set the tone for others in the workplace because your employees naturally look to you for leadership on a daily basis. By creating clear goals and expectations of others across the board, you create an environment of fairness. You cannot expect others to follow rules if you are not going to follow them yourself.
It is important not to give special or preferential treatment to certain employees or colleagues. Be aware of inherent, unconscious and implicit biases that may impact your interactions with others. Holding yourself and others accountable for their actions is important, but, again, it is not what you say but how you say it that is important to keep the environment positive.
Quash gossip in the workplace by not engaging in it. Discourage others from engaging in gossip, as well. Talking about others will make you less approachable, less trustworthy and will not promote positive relationships in the workplace.
It is not only important to pay attention to how we talk to others, but also how we talk to ourselves. Oftentimes, we are our own worst critic. Pay attention to your thoughts. Thinking that “things are never going to change,” “I can’t handle this” or “I don’t want to deal with this today” all are negative and defeating.
Combat negative thoughts by speaking to yourself positively. For example, “I can handle this,”
“Things are going to change; nothing every stays the same” and “This is an inconvenience, but I will handle it and then move on.” Don’t let negative thinking cause you to have a bad day.
Our self-talk habits have a much bigger effect on our mood and stress levels than we realize. When we verbalize positive statements for others to hear, we are promoting a positive workplace. Focus on what is going well rather than what is going wrong.
Find solutions to problems rather than complaining about them to others. Invite dialogue from others on how to solve problems. It’s okay not to always have the answers. It definitely is okay to ask for guidance when it is needed.
As a supervisor, you can promote positivity in the workplace by:
Engaged employees and coworkers are more productive and work better collaboratively. Employees who feel appreciated typically are more engaged. Engaged employees have better work performance and attendance. This will directly affect your work performance and stress levels, as well. Creating positivity in the workplace creates a win-win situation for management and employees.
Following the ideas presented in this article is a good start to promoting a healthy, positive work environment for everyone. If you find yourself struggling with having a positive attitude toward work, your Employee Assistance Program is here to help. For more information or assistance in promoting positivity in the workplace, visit EAP4YOU.com or contact your EAP at 800-327-4968
Categories: The Postal Supervisor
April 19, 2024
April 17, 2024

domain: sociology


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WISE Council approves $2 million in state funding for UL
With initial approval of state funds, the University of Louisiana at Lafayette is on a path to grow student enrollment and graduates in high demand workforce fields as outlined in the Workforce and Innovation for a Stronger Economy plan.
The WISE Council today approved the UL System’s campus plans for $9.5 million of state funding for workforce and research initiatives. An 83 percent in match funding brings the total investment to $17.4 million to be divided among UL System schools.
The UL System approved UL Lafayette’s plan for spending about $2 million in WISE funds last month. The university is on a path to receive a total investment of more than $3.3 million, with $1.3 million coming from private monies.
The UL System’s plan must still be approved by the Louisiana Board of Regents.
“This investment in our universities coupled with the 83 percent match funding will significantly help Louisiana close the workforce gaps,” said UL System President Sandra K. Woodley in a press release. “I am particularly proud of the high level of private industry support that will go towards new faculty and staff positions, scholarships, technology upgrades, and incentivizing research.”
Universities and colleges competed for shares of the $40 million, largely based on the degrees and certificates each produces in high-demand fields.
UL Lafayette administration plans to use its portion of WISE funds to provide more faculty and resources for students.
New hires would include two faculty positions and a recruitment and retention coordinator in School of Computing and Informatics, a petroleum engineering assistant professor and two instructors each for the engineering and nursing programs.
New scholarships would provide undergraduate and graduate stipends and assistantships to up to a dozen computer science students, a dozen engineering students and half a dozen nursing students. Instructional and research laboratories in engineering and computer science are also on the list to be enhanced.
“Our WISE plan, along with the financial support from the state and private sector, will help us to achieve greater success for our students and the state’s economy,” said Jim Henderson, provost and vice president of Academic Affairs at UL Lafayette, last month in a press release.
UL System WISE fund breakdown
Grambling State University
Louisiana Tech University
McNeese State University
Nicholls State University
Northwestern State University
Southeastern Louisiana University
University of Louisiana at Lafayette
University of Louisiana at Monroe
University of New Orleans
Source: UL System

domain: economics


*Atom platform for value market smartphones* what is new about this? It may be new for Intel, but in the market A9 based devices are already in every price slot you can imagine. what is the TDP and cost of one of these chips? what about the integrated graphics? One word - Disappointing.
@sylvie, thanks for the update. No doubt Intel failed to capture the mobile market just like the way ARM capture smartphone market but that doesn't mean we can ruleout Intel completely. The features which you have mentioned in your articles will definitely help Intel to recapture the lost market.
As we unveil EE Times’ 2015 Silicon 60 list, journalist & Silicon 60 researcher Peter Clarke hosts a conversation on startups in the electronics industry. Panelists Dan Armbrust (investment firm Silicon Catalyst), Andrew Kau (venture capital firm Walden International), and Stan Boland (successful serial entrepreneur, former CEO of Neul, Icera) join in the live debate.


Forget ‘Teenage Dream!’ Katy has a new hit on her hands, ‘Firework!’ Prepare to become obsessed.
I love, love, love Katy Perry‘s new song “Firework!” It’s super catchy and Katy’s voice sounds amazing, as always. But I have to admit… the new music video that hit the web Oct. 28 is pretty weird! Sparks literally burst first out of Katy’s chest, then out of a pregnant woman, and a child, who appears to be a cancer patient, looks on.
The whole thing is a bit strange, but hey, that’s Katy Perry!. What do YOU think of this new music video? Vote below!


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Disney is not letting the Frozen franchise go, with new voice actors in talks for the animated film's sequel.
Frozen 2, which has a November 27, 2019 release date in the United States, is looking to secure the voices of Evan Rachel Wood and Sterling K Brown.
The television stars are in talks to lend their voices according to Variety, which reports Idina Menzel, Kristen Bell and Josh Gad will reprise their roles from the 2013 hit film as Elsa, Anna, and Olaf, respectively.
The film will be directed by Jennifer Lee and Chris Buck, who led the first film, with Lee also scripting.
The original film made more than NZ$1.88 billion worldwide in 2013 and spawned a merchandising juggernaut, breaking numerous home video records.
It would go onto win both the Academy Award for Best Animated film and Best Original Song for 'Let It Go'. The writers of that song will also reportedly return for the sequel.
Most recently, Frozen skated its way onto the stage, with a Broadway production that was nominated for three Tony Awards.
Wood and Brown both received Emmy nominations this week for their roles on Westworld and This Is Us respectively, with Brown having previously won the television award for his star turn in American Crime Story: The People v. O. J. Simpson.

domain: artistic


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BBC Radio 1’s Live Lounge always brings good surprises. Electronic pop-duo AlunaGeorge stopped by to perform a silky smooth cover of Justin Bieber’s latest single “Company”. The cover features singer Aluna Francis on vocals over a stripped-down, minimal beat. The duo will be embarking on their solo tour with breakthrough “Gold” singer Kiiara this summer. In case you missed it, AlunaGeorge will also be joining Sia on her upcoming “Present is Nostalgic Tour” this fall. Exciting times for the talented pair. Check out AlunaGeorge’s cover of Justin Bieber’s “Company” below.
Currently browsing: AlunaGeorge
AlunaGeorge is back! And they’re back with a banger of a tune! “I’m In Control” is the British duo’s first single from their upcoming second album, titled “I Remember” and is the follow-up to 2013’s critically acclaimed “Body Music”. “I’m In Control” is a electro-pop song with the ever-so trendy tropical vibes and features some guest vocals rom Jamaican artist Popcaan. Together, they make quite the tune and gets us super excited for the new music to come! The single is out today on all digital retailers, via Island Records, so buy it now! For now, lose control to AlunaGeorge’s...more

domain: celebrity


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- 10 March 1884
- Chicago, Illinois, USA
Stuart Holmes, born Joseph Liebchen, was a silent screen leading man (from 1909) who starred in Fox's first feature film, Life's Shop Window (1914), filmed on Staten Island for $4,500. Being of somewhat menacing demeanour, the cold-eyed, moustachioed Holmes quickly discovered his penchant for playing dastardly villains of French, Italian or Russian extraction. He was highly rated by critics for his Grand Duke Michael in The Prisoner of Zenda (1922) and for Alexander, nemesis of Tess of the D'Urbervilles (1924). His characters rarely ever survived the final reel. After leaving Fox, Holmes joined Metro for similar work and then segued into character parts after the coming of sound. He was signed as a Warner Brothers extra in the mid-30's and continued to amass uncredited or cameo bits until his retirement in 1964, by which time he had appeared in some 530 films. His wife, Blanche Maynard, was a well-known Hollywood astrologer and Holmes himself (when not busy on screen) spent his free time wood-carving. He was reputedly rather good at it.
|Movie Name||Release Date|
|Ο άνθρωπος που γελά – The Man Who Laughs (1928)||November 4, 1928|
|Οι τέσσερις ιππότες της αποκάλυψης – The Four Horsemen of the Apocalypse (1921)||January 1, 1970|

domain: entertainment


The organ we had been servicing was poorly installed and was poorly laid out. The cramped Swell was positioned behind the Great and the façade pipes spoke poorly, functioning as the Great's 8' Diapason. We ultimately removed the instrument and started fresh in its chamber area. Rebuilding the organ with the Swell division on the first level with the Great division above it, gave us the necessary room for servicing and tuning access. All of the chests were rebuilt with electric valves and as is our standard, Plexiglas bottom boards. We added a new 8' Great Diapason leaving the façade but non-functioning. A new Zephyr blower was installed and the ICS-4000 was used as the control system.


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Videos por Caballero Portraits, Fotografos Expertos.
Caballero Jesse (c) 2016 Photographer
Hoy es nuestra quinceañera! Las fotos de nuestra fiesta y el video!
Hair and Make-up guru
Amazing Quinceanera pictures!
Jesse Caballero has been commissioned as the official photographer to document our wedding!
Caballero was the photographer chosen to document my quinceañera! I am so happy to be able to share the images and video with you!


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The Lajeunesse Royals know they can play a much better game than what they produced on Monday.
The No. 3-seeded Royals opened the OFSAA boys’ A basketball tournament with a 74-60 win over the No. 17-seed Oakville Ste. Trinite les Loups at the Lajeunesse gym.
“We were kind of not OFSAA ready mentally yet,” Lajeunesse’s Ali Aoude said. “It’s a different atmosphere with the crowd.
“Maybe after this first game, we can take some notes and lessons and get ready for the next games.”
The Royals built an 18-point lead in the second quarter, but got caught up in the attention of the home crowd and started to wildly throw up three-point attempts instead of running their offence.
“I think they were nervous and overly excited, especially with the whole school here,” Royals head coach Geoff Parent said of the gym crowd. “They start to play for the crowd and do things they don’t normally do.”
A 10-0 run by les Loups made things interesting, but Lajeunesse would never let the opposition get closer than eight points.
“It’s embarrassing,” Lajeunesse’s Ahmed El Hassan said. “We played horrible. I’m happy this was the first one.”
There were times in the game where the Royals turned up the defensive pressure and were rewarded with turnovers and transition baskets and also made Ste. Trinite pay for errant shots by pushing the ball up the floor.
“We have to push the ball more, play better defence, play more as a team and stay more positive,” Aoude said. “Out of this game, we have to take the good things out of the bad.”
Aoude led the Royals’ offence with 25 points while El Hassan added 14 points and Elio Skaf chipped in with 13.
“We can’t expect to play that way and move forward in this tournament,” Parent said. “If they can control their own emotions, which is the biggest battle, we should be OK.”
Next up for Lajeunesse is No. 8-ranked Kitchener Rockway Mennonite, which opened with a loss and then eliminated the No. 19-seeded Cardinal Carter Cougars from medal contention with a 76-57 win.
It was Cardinal Carter’s second loss of the day as the Cougars opened tournament play with an 85-56 loss to top-seeded Central Toronto Academy. Ryan Raffoul had 36 points for Cardinal Carter in the opener and added 29 points in the late game.

domain: game


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This DWS520KT is supplied with 2 x 1.5m guide rails, connector and TSTAK kit box.
The parallel plunge gives a smooth and constant hand position while cutting whilst the 48 tooth blade gives a fine and accurate finish and allows you to maintain a constant smooth cut. The saw has full wave electronics even under a heavy load it maintains a constant speed. The variable speed allows you to select the correct speed for different types of materials. The enclosed guard gives a 90% dust extraction capability when connected to a vacuum extractor. The saw’s 55mm depth of cut allows the trimming and sizing of most common door sizes. The 1.5m guide rails included in the kit feature marks on the rail so you can see exactly the start and end of where the blade will plunge. The anti-kickback function stops the saw from moving backwards along the rail and helps stop the saw from climbing out of the workpiece when making plunge cuts. Rail adjustment allows you to adjust the saw accurately to the rail. TSTAK case for the saw is supplied.
|Blade Diameter||165 mm|
|Bore Size||20 mm|
|Corded or Cordless||Corded|
|Motor Power||1,300 W|
|Nett Weight||5.1 kg|
|Sound Power Level [Uncertainty K]||103 dB(A) [3.0 dB]|
|Sound Pressure Level [Uncertainty K]||92 dB(A) [3.0 dB]|
|Spindle Lock||Standard Feature|
|Vibration Cutting Wood [Uncertainty K]||2.5 m/s² [1.5 m/s²]|
|Plug Fitted||UK 3 pin plug|
|Max Depth of Cut @ 45º||44 mm|
|Max Depth of Cut @ 90º||59 mm|
|Saw Blade Speed||1,750 - 4,200 rpm|

domain: hobby


Over the past few years, Analog Son has built a reputation throughout Colorado as being one of the tightest funk acts around. Since the band’s inception, co-founders Jordan Linit (guitar) and Josh Fairman (bass) have followed the basic tenets of the FUNK to a tee: freedom of expression; the fusion of jazz, soul, and R&B; and most importantly, filthy, filthy grooves to put the stank on that face. The group is set to headline Boulder, Colorado’s The Fox Theatre on Thursday, October 12th, with support from PHO and Envy Alo (purchase tickets here).With two proper studio albums under their belt including the recent 2017 release of Black Diamond, the formidable duo behind Analog Son has been joined by the likes of New Orleans royalty Ivan Neville, singer Corey Frye (The Main Squeeze), Nigel Hall (Lettuce) percussionist Jason Hann (The String Cheese Incident), keyboardist Joe Tatton (The New Mastersounds), both Ryan Zoidis and Eric “Benny” Bloom of The Shady Horns, and many more in the studio. Given that Fairman—who also recently joined fellow Colorado act SunSquabi—has his own recording studio where he has produced many of the aforementioned artists and then some, Analog Son’s studio albums are consistently all-star efforts worthy of a listen. In the live setting, Analog Son taps a loose collection of artists to join them on stage and help create the high-energy funk shows that they’ve become known for.Hailing from the Twin Cities (Minneapolis, MN, FYI), instrumental funk act PHO blends modern and classic influences to take their music beyond all musical boundaries. The group has toured around the country with The Motet, Dopapod, and Jonathan Richman, to name a few. Their biggest “WTF! Is this really happening?” moment came when Prince himself found the band on Youtube and posted about them, subsequently inviting them to play at Paisley Park in support of Larry Graham. If The Purple One’s personal recommendation isn’t enough, then you have to ask yourself, what is?Old-school organ funk and progressive guitar rock drive the “boogaloo fusion” sound that defines Boulder-based trio Envy Alo. With their immense musicality at the forefront, the trio consisting of Aaron Pettine (synth/organ), Kevin Hinder (guitar), and Nate Etter (drums) has steadily been gaining a following across Colorado since their inception in 2015. The band had a standout performance in Boulder back in July of this year, which truly put this young group on the map. This time around, expect the group to go even bigger for their upcoming return to the Fox Theatre.Analog Son guitarist Jordan Linit discussed his upcoming show with Analog Son at the Fox Theatre, which follows a summer that witnessed some big moments for the band, “Analog Son has been sounding better than ever. We’re coming off an extreme high after our first Red Rocks show with The String Cheese Incident this summer. The Fox Theater is a gem in one of the nation’s most vibrant music scenes. Come watch us tear the roof off!”Tickets for Analog Son with PHO and Envy Alo on Thursday, Oct. 12th are currently on-sale and can be purchased here. For show updates and additional information, join the Facebook Event page.[cover photo courtesy of Analog Son’s FB page and J. Mimna Photography]Enter To Win A Pair Of Tickets Below!
A recently retired Tipperary hurler says there’s now a marked difference in the fitness levels of intercounty and club players.Paddy Stapleton – who stepped away from the county panel last month – believes this is down to the increased amount of gym work that the former group are doing.The Borrisoleigh club man was involved with the Tipp set-up for 10 years. He says the physical demands put on elite players changed hugely in that time