We thank M.
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Maudsley, M. Weathers and members of the Altizer lab for assistance capturing fall migrants. We also appreciate helpful comments from two anonymous reviewers of the manuscript. National Center for Biotechnology Information , U. PLoS One. Published online Apr 2. Haley Johnson , 1 Michelle J.
Solensky , 1 Dara A. Satterfield , 2 and Andrew K. Michelle J. Dara A. Andrew K. Guy Smagghe, Editor. Author information Article notes Copyright and License information Disclaimer. Competing Interests: The authors have declared that no competing interests exist. Received Dec 17; Accepted Mar 6. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.
Associated Data Supplementary Materials File S1: Summary of additional measurements leg size of monarch specimens, which were obtained to help interpret results concerning wing size, as well as further analyses that examined leg size variation among treatment groups. Abstract In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage.
Introduction In animals with biphasic life cycles, the conditions experienced by larvae can greatly affect the fitness of adults  ,  ,  , . Open in a separate window. Figure 1. Male monarch butterfly Danaus plexippus , showing the distinctive orange and black wing patterns characteristic of this species. Photograph from Wikimedia commons. Materials and Methods Ethics statement No permits were necessary for the collection of butterflies or for the experimental work presented here, and this project did not involve endangered or protected species.
Butterflies Before the experiment began, we first created a captive collection of adult monarchs to use as breeding stock.
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Collecting migrants Based on specific outcomes of the food experiment regarding wing morphology, we also examined wing scans from several previously-collected sets of migrant monarchs  ,  to evaluate whether certain patterns observed in the captive monarchs represented what would be seen in free-living monarchs.
Data analysis The response variables for the food stress experiment included monarch forewing size area , shape aspect ratio , orange hue, and the two measures of melanism percent black and black density. Figure 2. Scanned image of a monarch butterfly top panel with one forewing cropped to show details of forewing measurements. Figure 3. Depiction of forewing color measurements top left , including orange hue and black density scores.
Table 1 Relationships between all response variables in the food stress experiment for both male and female monarchs. Pearson correlation coefficients are shown for each pairwise combination. For the black density score, smaller values indicate darker pigment. Table 2 Summary of univariate results for each response variable wing morphology traits of adult monarchs in this study. See text for descriptions of variables. Food treatments were control food given ad libitum to larval monarchs , low stress foodplant removed for 24 hours at the 4 th instar and high stress foodplant removed for 24 hours at both the 4 th and 5 th instar.
Figure 4. Plots showing the effect of larval food treatment on adult monarch wing size A and shape B. Figure 5. Plots showing the effect of larval food treatment on adult monarch wing color, including orange hue A , proportion of black B and the intensity of black pigment C. Figure 6. Relationship between monarch wing melanism and shape in lab-reared captive, A and wild specimens captured during fall migration B. Note that the X and Y axes of both plots differ. Discussion The overarching goal of this study was to determine the effects, if any, of larval food supply on adult wing morphology in monarch butterflies.
Supporting Information File S1 Summary of additional measurements leg size of monarch specimens, which were obtained to help interpret results concerning wing size, as well as further analyses that examined leg size variation among treatment groups. DOCX Click here for additional data file. Acknowledgments We thank M. Funding Statement The authors have no support or funding to report.
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References 1. Hahn DA Larval nutrition affects lipid storage and growth, but not protein or carbohydrate storage in newly eclosed adults of the grasshopper Schistocerca americana. Journal of Insect Physiology 51 : — Scott DE The effect of larval density on adult demographic traits in Ambystoma opacum. Ecology 75 : — Telang A, Wells MA The effect of larval and adult nutrition on successful autogenous egg production by a mosquito.
Journal of Insect Physiology 50 : — Tigreros N Linking nutrition and sexual selection across life stages in a model butterfly system. Functional Ecology 27 : — Annual Review of Entomology 47 : — Ecological Entomology 34 : — Oecologia : — Brower LP Understanding and misunderstanding the migration of the monarch butterfly Nymphalidae in North America: — Journal of the Lepidopterists' Society 49 : — In: Oberhauser K, Solensky M, editors.
The monarch butterfly Biology and Conservation. Altizer S, Davis AK Populations of monarch butterflies with different migratory behaviors show divergence in wing morphology. Evolution 64 : — Journal of Thermal Biology 30 : — Insects 4 : — PloS One 7 : e doi: Behavioral Ecology 24 : — Dmitriew C, Rowe L The effects of larval nutrition on reproductive performance in a food-limited adult environment.
Plos One 6 : 6. Kemp DJ Resource-mediated condition dependence in sexually dichromatic butterfly wing coloration. Evolution 62 : — Plos One 7 : 8. Knuttel H, Fiedler K Host-plant derived variation in ultraviolet wing pattern influences mate selection by male butterflies.
Journal of Experimental Biology : — Evolution 58 : — Pleasants JM, Oberhauser KS Milkweed loss in agricultural fields because of herbicide use: effect on the monarch butterfly population. Insect Conservation and Diversity: Online Early doi: Altizer SM, Oberhauser K, Brower LP Associations between host migration and the prevalence of a protozoan parasite in natural populations of adult monarch butterflies. Ecological Entomology 25 : — The review must be at least 50 characters long. The title should be at least 4 characters long. Your display name should be at least 2 characters long.
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Close Report a review At Kobo, we try to ensure that published reviews do not contain rude or profane language, spoilers, or any of our reviewer's personal information. Would you like us to take another look at this review? No, cancel Yes, report it Thanks! Eleuthero supplementation also seemed to increase FFA availability by increasing fat oxidation, sparing muscle glycogen. They concluded eleuthero may be an effective nutritional supplement for endurance athletes, but the exact mechanisms involved need further investigation.
The only side effect reported was mild insomnia in one participant for the first day of eleuthero supplementation. The authors reported no financial interest in eleuthero as a supplement. Cordyceps sinensis is a naturally occurring parasitic fungus native to the Tibetan Plateau. Although wild Cordyceps increasingly is rare in its natural habitat and thus very expensive, a refined standardized fermentation product, Cs-4, is produced from mycelial strains isolated from wild Cordyceps and contains similar chemical constituents 0.
Researchers from UCLA conducted a double-blind, placebo-controlled, prospective study of the commercially available Cs-4 product on 20 men and women between 50 and 75 years of age. Subjects were randomized to Cs-4 or placebo starch capsules, taken three times a day with water or food for 12 weeks. Exercise performance was evaluated on a cycle ergometer, while physical fitness was assessed through measurements of VO 2max and gas exchange metabolic threshold. Dropouts in each group were similar at study completion, with eight in the Cs-4 group and seven in the placebo group.
From baseline, the Cs-4 group had an increase in metabolic threshold of There were no other significant differences in VO 2max , heart rate, maximum work rate or maximum ventilation, blood pressure, lactic acid peak level, recovery, or heart rate recovery. The authors believed this small but consistent improvement in aerobic performance of subjects receiving Cs-4 would translate to meaningful improvements in activities of daily living in older adults if it could be reproduced in larger studies. Like cordyceps, Rhodiola crenulata , which is closely related to Rhodiola rosea and grows mostly in the Tibetan region, is another botanical native to cold, mountainous regions of Asia, Europe, and high altitudes in the Arctic.
It also has a long history of use in traditional Asian medical systems for a wide variety of conditions, including improved physical endurance and work performance, resistance to high-altitude sickness, mental capacity, and longevity.
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Today, rhodiola most commonly is used for increasing energy, endurance, strength, and mental capacity. Chen et al conducted a double-blind, placebo-controlled trial on 18 male long-distance track and field athletes during high-altitude training 2, meters. The groups participated in intensive exercise training, which included distance, speed, endurance, and strength workouts.
There were no differences in VO 2max , red blood cell, hemoglobin, testosterone, or cortisol levels. The authors concluded that RC improves aerobic exercise capacity and produces anti-stress reactions that may increase adaptations to exercise training at high altitude. Ashwagandha Withania somnifera , also known as Indian ginseng or winter cherry, is a dense shrub with roots that have been used extensively in Ayurvedic medicine for 3, years as a general tonic to help the body adapt to stress.
The root contains a number of antioxidants as well as flavonoids, alkaloids, and steroidal lactones, which are thought to confer the beneficial effects in the body. A randomized, double-blind, prospective clinical study of 57 male subjects 18 to 50 years of age with minimal experience in resistance training were divided into treatment and placebo groups. Participants agreed to refrain from tobacco, alcohol, and anti-inflammatory agents during the study and had to be cleared for participation by their physician. The treatment group reviewed mg Ashwagandha root twice daily, while the control received a starch placebo.
Muscle strength, muscle size, body composition, serum testosterone level, and muscle recovery was measured at baseline and then again after eight weeks of a resistance training program. The three-days-weekly training program targeted large muscle groups with instruction on proper technique. Muscle strength was measured using the one repetition maximum for bench press and leg extension exercise. Serum creatine kinase was used as a marker of muscle injury to evaluate recovery.
Larva High School 2 - Expectations and Endurance
The treatment group had significantly greater improvement in muscle strength in both exercises Ashwagandha 1, Ashwagandha: The authors concluded that Ashwagandha may be useful to take in a resistance training program. Shenoy et al studied supplementation of Ashwagandha in 40 elite cyclists 20 of each gender , defined as participation of the athlete in at least state-level events. They were divided into treatment mg capsules of standardized Ashwagandha root extract twice daily and placebo starch tablets.
A baseline graded-exercise test GXT was performed on a treadmill with electronic heart rate monitor and full nose-mouth piece to measure VO 2max , respiratory exchange ratio RER , and total time for the athlete to reach their volitional point of exhaustion.
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