, Volume 24, Issue 5, pp 865–877 | Cite as

Seasonal variation in daily insolation induces synchronous bud break and flowering in the tropics

  • Zoraida Calle
  • Boris O. Schlumpberger
  • Lorena Piedrahita
  • Avigdor Leftin
  • Steven A. Hammer
  • Alan Tye
  • Rolf BorchertEmail author
Original Paper


In many temperate plants seasonal variation in day length induces flowering at species-specific times each year. Here we report synchronous bud break and flowering of tropical perennials that cannot be explained by seasonal changes in day length. We recorded flushing and flowering of more than 100 tropical trees, succulents and understory herbs over several years. We observed the following phenological patterns throughout the northern Neotropics: wide-ranging trees flush or flower twice a year at the Equator, but annually further north; many trees leaf out in February; in autumn, wide-ranging perennials flower 4 months earlier in Mexico than at the Equator. This latitudinal variation of phenology parallels that of the annual cycle of daily insolation, a function of day length and solar irradiation. Insolation has two annual maxima at the Equator, it rapidly increases in February at all latitudes, and between Mexico and the Equator its maximum shifts from the summer solstice to the autumn equinox. These unique, manifold correlations suggest that throughout the tropics insolation, rather than day length, may control the phenology of many perennials. Our observations significantly extend current knowledge of environmental signals involved in photoperiodic control of plant development.


Day length Phenology Photoperiod Solar irradiation Succulents Tropical trees 



A. H. Strahler (Boston University) gave valuable advice on insolation. R. Guy (University of British Columbia), U. Lüttge (Technical University Darmstadt), S. S. Renner (University of Munich) and three anonymous reviewers made valuable suggestions for improving the manuscript. E. V. Jaarsveld (Cape Town) shared observations of the flowering phenology of Southwest African succulents. F. Berger supported the observation of succulent phenology in Munich, and M. Uhlig (Uhlig Kakteen) provided numerous Rebutia fiebrigii. G. Pakkad and Thonglaw Srithong (Forest Restoration Unit, Chiang Mai University, Chiang Mai, Thailand) guided RB to trees flushing on February 15, 2010.

Supplementary material

468_2010_456_MOESM1_ESM.doc (316 kb)
Supplementary material 1 (DOC 315 kb)


  1. Black GA, Dobzhansky T, Pavan C (1950) Some attempts to estimate species diversity and population density of trees in Amazonian forests. Bot Gaz 111:413–425CrossRefGoogle Scholar
  2. Böhlenius H, Huang T, Charbonnel-Campaa L, Brunner AM, Jansson S, Strauss S, Nilsson O (2006) CO/FT module controls timing of flowering and seasonal growth cessation in trees. Science 312:1040–1043CrossRefPubMedGoogle Scholar
  3. Borchert R (1994a) Soil and stem water storage determine phenology and distribution of tropical dry forest trees. Ecology 75:1437–1449CrossRefGoogle Scholar
  4. Borchert R (1994b) Induction of rehydration and bud break by irrigation or rain in deciduous trees of a tropical dry forest in Costa Rica. Trees 8:198–204CrossRefGoogle Scholar
  5. Borchert R (1996) Phenology and flowering phenology of neotropical dry forest species: evidence from herbarium collections. J Trop Ecol 12:65–80CrossRefGoogle Scholar
  6. Borchert R, Rivera G (2001) Photoperiodic control of seasonal development and dormancy in tropical stem-succulent trees. Tree Physiol 21:213–221PubMedGoogle Scholar
  7. Borchert R, Renner SS, Calle Z, Navarrete D, Tye A, Gautier L, Spichiger R, von Hildebrand P (2005a) Photoperiodic induction of synchronous flowering near the Equator. Nature 433:627–629CrossRefPubMedGoogle Scholar
  8. Borchert R, Robertson K, Schwartz MD, Williams-Linera G (2005b) Phenology of temperate trees in tropical climates. Int J Biometeorol 50:57–65CrossRefPubMedGoogle Scholar
  9. Bullock SH (2002) La fenologia de plantas en Chamela. In: Noguera FA, Vega-Rivera JG, Garcia-Aldrete AN, Quesada-Avendano M (eds) Historia Natural de Chamela. Instituto de Biología, UNAM, Mexico, pp 491–498Google Scholar
  10. Calle Z, Strahler AH, Borchert R (2009) Daily insolation induces synchronous flowering flowering of Montanoa and Simsia (Asteraceae) between Mexico and the Equator. Trees 23:1247–1254CrossRefGoogle Scholar
  11. Cleland EE, Chuine I, Menzel A, Mooney HA, Schwarz MD (2007) Shifting plant phenology in response to global change. Trends Ecol Evol 22:357–365CrossRefPubMedGoogle Scholar
  12. Croat TB (1969) Seasonal flowering behavior in central Panama. Ann Missouri Bot Gard 56:295–307Google Scholar
  13. Dornelas MC, Rodriguez APM (2005) The rubber tree (Hevea brasiliensis Muell.Arg.) homologue of the LEAFY/FLORICAULA gene is preferentially expressed in both male and female floral meristems. J Exp Bot 56:1965–1974CrossRefPubMedGoogle Scholar
  14. Downs RJ, Borthwick HA (1956) Effects of photoperiod upon the growth of trees. Bot Gaz 117:310–326CrossRefGoogle Scholar
  15. Eamus D (1999) Ecophysiological traits of deciduous and evergreen woody species in the seasonally dry tropics. Trends Ecol Evol 14:11–16CrossRefPubMedGoogle Scholar
  16. Elliott S, Baker PJ, Borchert R (2006) Leaf flushing during the dry season: the paradox of Asian monsoon forests. Global Ecol Biogeogr 15:248–257Google Scholar
  17. Funk VA (1982) The systematics of Montanoa (Asteraceae, Heliantheae). Mem N Y Bot Gard 36:1–133Google Scholar
  18. Garner WW, Allard HA (1920) Effect of the relative length of day and night and other factors of the environmental on growth and reproduction of plants. J Agric Res 18:553–606Google Scholar
  19. Heide O (1974) Growth and dormancy in Norway spruce ecotypes (Picea abies). Physiol Plant 30:1–12CrossRefGoogle Scholar
  20. Holttum RE (1940) Periodic leaf-exchange and flowering of trees in Singapore (II). Gardens Bull S S 11:119–175Google Scholar
  21. Hubbell SP, He F, Condit R, Borda-de-Agua L, Kellner J, ter Steege H (2008) How many tree species are there in the Amazon and how many will go extinct? PNAS 105:11498CrossRefPubMedGoogle Scholar
  22. Jackson SD (2008) Plant responses to photoperiod. New Phytol 181:517–531CrossRefGoogle Scholar
  23. Janzen DH (1970) Jacquinia pungens, a heliophile from the understory of tropical deciduous forests. Biotropica 2:112–119CrossRefGoogle Scholar
  24. Janzen DH (1982) Natural history of guacimo fruits (Sterculiaceae, Guazuma ulmifolia) with respect to consumption by large mammals. Am J Bot 69:1240–1250CrossRefGoogle Scholar
  25. Jian M, Liu Q, Tang P, Liang Y (2009) Floristic analysis of the evergreen broad-leaved forest community from Jiulianshan National Reserve in Jiangxi Province, China. Front For China 4:416–423CrossRefGoogle Scholar
  26. Juntilla O (1980) Effect of photoperiod and temperature on apical growth cessation in two ecotypes of Salix and Betula. Physiol Plant 48:347–352CrossRefGoogle Scholar
  27. Kira T (1991) Forest systems of East and South East Asia in a global perspective. Ecol Res 6:185–200CrossRefGoogle Scholar
  28. Kőrner C, Basler D (2010) Phenology under global warming. Science 327:1461–1462CrossRefPubMedGoogle Scholar
  29. Kramer PJ (1936) Effect of variation in length of day on growth and dormancy of trees. Plant Physiol 11:127–137CrossRefPubMedGoogle Scholar
  30. Kushwara CP, Singh KP (2005) Diversity of leaf phenology in a tropical deciduous forest in India. J Trop Ecol 21:47–56CrossRefGoogle Scholar
  31. Lüttge U, Hertel B (2009) Diurnal and annual rhythms of trees and their relations to the circadian clock. Trees 23:683–700CrossRefGoogle Scholar
  32. Melzer S, Lens F, Gennen J, Vanneste S, Rohde A, Beeckman T (2008) Flowering-time genes modulate meristem determinacy and growth form in Arabidopsis thaliana. Nat Genet 40:1489–1492Google Scholar
  33. Menzel A, Fabian P (1999) Growing season extended in Europe. Nature 397:659CrossRefGoogle Scholar
  34. Newstrom LE, Frankie GW, Baker HG (1994) A new classification for plant phenology based on flowering patterns in lowland tropical rain forest trees at la Selva, Costa Rica. Biotropica 26:141–159CrossRefGoogle Scholar
  35. Njoku E (1963) Seasonal periodicity in the growth and development of some forest trees in Nigeria. I. Observations on mature trees. J Ecol 51:617–624CrossRefGoogle Scholar
  36. Opler PA, Frankie GW, Baker HG (1976) Rainfall as a factor in the release, timing and synchronization of anthesis by tropical trees and shrubs. J Biogeogr 3:221–236CrossRefGoogle Scholar
  37. Paredes-Aguilar R, van Devender TR, Felger RS (2000) Cactáceas de Sonora, Mexico, su diversidad, uso y conservación. Arizona Desert Museum, TucsonGoogle Scholar
  38. Perry TO, Wang CW (1960) Genetic variation in winter chilling requirements for the date of dormancy break in Acer rubrum. Ecology 41:790–794CrossRefGoogle Scholar
  39. Pőhlmann G (1976) Atlas der Volksrepublik China. TFH Berlin, BerlinGoogle Scholar
  40. Primack D, Imbres C, Primack RB, Miller-Rushing AJ, Del Tredici P (2004) Herbarium specimens demonstrate earlier flowering in response to warming in Boston. Am J Bot 91:1260–1264CrossRefGoogle Scholar
  41. Ray PM, Alexander WE (1966) Photoperiodic adaptation to latitude in Xanthium strumarium. Am J Bot 53:806–816CrossRefGoogle Scholar
  42. Reich PB, Borchert R (1982) Phenology and ecophysiology of the tropical tree, Tabebuia neochrysantha (Bignoniaceae). Ecology 63:294–299CrossRefGoogle Scholar
  43. Renner SS (2007) Synchronous flowering linked to changes in solar radiation intensity. New Phytol 175:195–197CrossRefPubMedGoogle Scholar
  44. Rivera G, Borchert R (2001) Induction of flowering in tropical trees by a 30-min reduction in photoperiod: evidence from field observations and herbarium specimens. Tree Physiol 21:202–212Google Scholar
  45. Rivera J, Cozza J (2008) Reduced photoperiod induces partially synchronous flowering in an understory rain forest herb, Begonia urophylla, in Costa Rica. Biotropica 40:363–365CrossRefGoogle Scholar
  46. Rivera G, Elliott S, Caldas LS, Nicolossi G, Coradin VTR, Borchert R (2002) Increasing day length induces spring flushing of tropical trees in the absence of rain. Trees 16:445–456CrossRefGoogle Scholar
  47. Smith GF, Chesselet P, Jaarsveld EV, Hartmann HEK, Hammer S, van Wyk B-E, Burgoyne P, Klak C, Kurzweil H (1998) Mesembs of the world. Briza Publications, PretoriaGoogle Scholar
  48. Strahler AN, Strahler A (2005) Introducing physical geography. Wiley, New YorkGoogle Scholar
  49. Thomas B, Vince-Prue D (1997) Photoperiodism in plants. Academic Press, San DiegoGoogle Scholar
  50. Valiente-Banuet A, Arizmendi M, Rojas-Martinez A, Dominguez-Canseco L (1995) Ecological relationships between columnar cacti and nectar-feeding bats in Mexico. J Trop Ecol 12:103–119CrossRefGoogle Scholar
  51. Van Schaik CP, Terborgh JW, Wright SJ (1993) The phenology of tropical forests, adaptive significance and consequences for primary consumers. Annu Rev Ecol Syst 24:353–377CrossRefGoogle Scholar
  52. Von Willert DJ, Eller EM, Werger MJA, Brickmann E, Ihlenfeld H-D (1992) Life strategies of succulents in deserts. Cambridge University Press, CambridgeGoogle Scholar
  53. Wareing PF (1956) Photoperiodism in woody plants. Ann Rev Plant Pysiol 7:191–214CrossRefGoogle Scholar
  54. Williams RJ, Myers BA, Muller WA, Duff GA, Eamus D (1997) Leaf phenology of woody species in a Northern Australian tropical savanna. Ecology 78:2542–2558CrossRefGoogle Scholar
  55. Williams-Linera G (1997) Phenology of deciduous and broadleaved tree species in a Mexican tropical lower montane forest. Global Ecol Biogeogr Lett 6:115–127CrossRefGoogle Scholar
  56. Windsor D (1982) The phenology of selected trees at Barro Colorado Island, Republic of Panama. Smithsonian Tropical Research Institute, Panama, p 189Google Scholar
  57. Wright SJ (1996) Phenological responses to seasonality in tropical forest plants. In: Mulkey SS, Chazdon RL, Smith AP (eds) Tropical forest ecophysiology. Chapman and Hall, New York, pp 440–460Google Scholar
  58. Yeang H-Y (2007a) Synchronous flowering of the rubber tree (Hevea brasiliensis) induced by high solar intensity. New Phytol 175:283–289CrossRefPubMedGoogle Scholar
  59. Yeang H-Y (2007b) The sunshine-mediated trigger of synchronous flowering in the tropics: the rubber tree as a study model. New Phytol 176:730–735CrossRefPubMedGoogle Scholar
  60. Zimmerman JK, Wright SJ, Calderon O, Aponte Pagan M, Paton SR (2007) Flowering and fruiting phenologies of seasonal and aseasonal neotropical forests: the role of annual changes in irradiance. J Trop Ecol 23:231–251CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2010

Authors and Affiliations

  • Zoraida Calle
    • 1
  • Boris O. Schlumpberger
    • 2
    • 7
  • Lorena Piedrahita
    • 1
  • Avigdor Leftin
    • 3
  • Steven A. Hammer
    • 4
  • Alan Tye
    • 5
  • Rolf Borchert
    • 6
    Email author
  1. 1.Center for Research on Sustainable Agriculture CIPAVCaliColombia
  2. 2.Department of BiologyUniversity of MunichMunichGermany
  3. 3.Department of ChemistryUniversity of ArizonaTucsonUSA
  4. 4.Sphaeroid InstituteVistaUSA
  5. 5.Secretariat of the Pacific Regional Environment ProgrammeApiaSamoa
  6. 6.Division of Biological SciencesUniversity of KansasLawrenceUSA
  7. 7.Herrenhausen GardensHannoverGermany

Personalised recommendations