We hypothesized differences in molecular strategies for similar journeys that migrants undertake to reproduce in spring and to overwinter in autumn. We tested this in redheaded buntings () photoinduced into spring and autumn migratory states, with winter and summer non-migratory states as controls. Compared with controls, buntings fattened, gained weight and showed (nocturnal migratory restlessness) in the migratory state. Spring migration was associated with greater fat and body mass, and higher intensity of, compared with autumn migration. Circulating corticosterone levels were higher in spring, while T3 levels were higher in autumn. Hypothalamic expression of thyroid hormone-responsive (, ), light-responsive (, ,) and (tyrosine hydroxylase, involved in dopamine biosynthesis) genes showed significant changes with transition from non-migratory to the migratory state. There were significantly higher mRNA expressions in autumn, except for higher levels in the spring. Furthermore, the expression patterns of (not) and genes suggested an epigenetic difference between the non-migrant and migrant periods, and the spring and autumn migrant periods. These results demonstrate for the first time seasonal transition in hypothalamic gene expressions, and suggest differences in regulatory strategies at the transcriptional level for spring and autumn migrations in songbirds.
The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4Wm(-2). The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D= 6:6:1:11 h) at 0.25+/-0.01 W m(-2); two light periods applied were of the same (450nm: blue:blue, B:B; 640nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LLdim; approximately 0.01 Wm(-2), the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LLdim, all birds free-ran with a period >24h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.
This study analyzed the role of day length in regulation of seasonal body fattening and testicular growth in a latitudinal Palaearctic-Indian migrant, the redheaded bunting (Emberiza bruniceps). When exposed to increasing photoperiods (hours of light: hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D, 13L:11D, 14L:10D, and 18L:6D) for 9-12 weeks, buntings responded in a photoperiod-dependent manner and underwent growth and regression cycle under photoperiods of > or =12 hr per day. Also, the response to a long photoperiod of birds that were held under natural photoperiods at 27 degrees N for 2 years was similar to those who arrived the same year from their breeding grounds ( approximately 40 degrees N), suggesting that the experience of higher amplitude day-night (light-dark, LD) cycles during migratory and breeding seasons were not critical for the subsequent response (initiation-termination-reinitiation) cycle. Another experiment examined entrainment of the circadian photoperiodic rhythm in buntings by subjecting them to T=24+/-2 hr LD-cycles with 8 hr photophase and to T=22 and 24 hr with 11 hr photophase. The results showed a reduction in critical day length under T=22 hr LD-cycle. In the last experiment, we constructed an action spectrum for photoperiodic induction by exposing birds for 4.5 weeks to 13L:11D of white (control), blue (450 nm), or red (640 nm) light at irradiances ranging from 0.028 to 1.4 W m(-2). The threshold light irradiance for photoinduction was about 10-fold higher for blue light, than for red and white lights. These results conclude that the daily light of the environment regulates the endogenous program that times seasonal responses in body fattening and testicular cycles of the redheaded bunting.
Innovation is fundamental to the process of an economy’s growth and is crucial for its survival in today’s dynamic world. Investing in innovation, therefore, is a requisite for developing and developed countries alike. The innovation revolution has been as manifesting as the Industrial Revolution. Yet, its patterns are unevenly distributed. While some regions have witnessed a phenomenal growth in the level of patenting activity, other regions have shown a decline in the innovative output. With this backdrop, the present article analyses various institutional and macroeconomic variables that impact the level of innovation for a set of 15 Asian countries. Using two-step system generalized method of moments (GMM) technique on a panel dataset for the period 2008–2017, the study examines the determinants of macroeconomic indicators on the level of innovation (as measured by the country’s patenting activity). The econometric results indicate that institutional quality, education and trade openness influence innovation favourably. Further, foreign direct investments have a negative impact on the level of innovation. A key finding of emerging from the analysis is the U-shaped relationship between financial development and innovation. Based on the results, the article concludes from a broad policy perspective.
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