, Volume 147, Issue 3, pp 396–405 | Cite as

Tuber size variation and organ preformation constrain growth responses of a spring geophyte

  • Marinus J. A. WergerEmail author
  • Heidrun Huber


Functional responses to environmental variation do not only depend on the genetic potential of a species to express different trait values, but can also be limited by characteristics, such as the timing of organ (pre-) formation, aboveground longevity or the presence of a storage organ. In this experiment we tested to what degree variation in tuber size and organ preformation constrain the responsiveness to environmental quality and whether responsiveness is modified by the availability of stored resources by exposing the spring geophyte Bunium bulbocastanum to different light and nutrient regimes. Growth and biomass partitioning were affected by initial tuber size and resource availability. On average, tuber weight amounted to 60%, but never less than 30% of the total plant biomass. Initial tuber size, considered an estimate of the total carbon pool available at the onset of treatments, affected plant growth and reproduction throughout the experiment but had little effect on the responsiveness of plants to the treatments. The responsiveness was partly constrained by organ preformation: in the second year variation of leaf number was considerably larger than in the first year of the treatments. The results indicate that a spring geophyte with organ preformation has only limited possibilities to respond to short-term fluctuations of the environment, as all leaves and the inflorescence are preformed in the previous growing season and resources stored in tubers are predominantly used for survival during dormancy and are not invested into plastic adjustments to environmental quality. Such spring geophytes have only limited possibilities to buffer environmental variation. This explains their restriction to habitats characterized by predictable changes of the environmental conditions.


Biomass partitioning Life history strategy Light Nutrient Organ preformation Storage Tuber size 



We thank Fan Zhang for invaluable help during the experiment, and Heinjo During and Josef Stuefer and two anonymous referees for insightful comments on an earlier version of this manuscript.


  1. Abrahamson WG (1979) Patterns of resource allocation in wildflower populations of fields and woods. Am J Bot 66:71–79CrossRefGoogle Scholar
  2. Aydelotte AR, Diggle PK (1997) Analysis of developmental preformation in the alpine herb Caltha leptosepala. Am J Bot 84:1646–1657CrossRefGoogle Scholar
  3. Beard JS (1983) Ecological control of the vegetation of southwestern Australia: moisture versus nutrients. In: Kruger FJ, et al (eds) Mediterranean-type ecosystems. Springer, Berlin Heidelberg New York, pp 66–73Google Scholar
  4. Billings WD, Mooney HA (1968) The ecology of arctic and alpine plants. Biol Rev 43:481–529CrossRefGoogle Scholar
  5. Bobbink R, Willems JH (1987) Increasing dominance of Brachypodium pinnatum (L.) Beauv. in chalk grasslands: a threat to a species-rich ecosystem. Biol Cons 40:301–314CrossRefGoogle Scholar
  6. Chapin FS, Schulze ED, Mooney HA (1990) The ecology and economics of storage in plants. Annu Rev Ecol Syst 21:423–477CrossRefGoogle Scholar
  7. Diggle PK (1997) Extreme preformation in alpine Polygnonum viviparum: An architectural and developmental analysis. Am J Bot 84:154–169CrossRefGoogle Scholar
  8. Ehrlen J, van Groenendael J (2001) Storage and the delayed costs of reproduction in the understorey perennial Lathyrus vernus. J Ecol 89:237–246CrossRefGoogle Scholar
  9. Elemans M (2005) Plant traits in forest understory herbs—a modeling study. PhD thesis, Utrecht UniversityGoogle Scholar
  10. Geber MA, de Kroon H, Watson MA (1997a) Organ preformation in mayapple as a mechanism for historical effects on demography. J Ecol 85:211–223CrossRefGoogle Scholar
  11. Geber MA, Watson MA, de Kroon H (1997b) Organ preformation, development and resource allocation in perennials. In: Bazzaz FA, Grace J (eds) Plant resource allocation. Academic, New York, pp 113–141CrossRefGoogle Scholar
  12. Harper JL (1977) Population biology of plants. Academic, London Google Scholar
  13. Hegi G (1975) Illustrierte Flora von Mitteleuropa. Band V/2. P.Parey, BerlinGoogle Scholar
  14. Huber H, Stuefer JF, Willems JH (1996) Environmentally induced carry-over effects on seed production, germination and seedling performance in Bunium bulbocastanum (Apiaceae). Flora 191:353–361Google Scholar
  15. Huber H, Fijan A, During HJ (1998) A comparative study of spacer plasticity in erect and stoloniferous herbs. Oikos 81:576–586CrossRefGoogle Scholar
  16. Huber H, Whigham DF, O’Neill J (2004) Time of disturbance can change life history decisions in the clonal forest understory herb Uvularia perfoliata. Evol Ecol 18:521–539CrossRefGoogle Scholar
  17. Inouye DW (1986) Long-term preformation of leaves and inflorescences by a long-lived perennial monocarp, Frasera speciosa (Gentianaceae). Am J Bot 73:1535–1540CrossRefGoogle Scholar
  18. Irmisch T (1854) Beiträge zur vergleichenden Morphologie der Pflanzen I. Abhandlungen der naturforschenden Gesellschaft zu Halle 2:31–80Google Scholar
  19. Iwasa Y, Cohen D (1989) Optimal growth schedule of a perennial plant. Am Nat 133:480–505CrossRefGoogle Scholar
  20. Iwasa Y, Kubo T (1997) Optimal storage for recovery after unpredictable disturbances. Evol Ecol 11:41–65CrossRefGoogle Scholar
  21. Kawano S (1985) Life history characteristics of temperate woodland plants in Japan. In: White J (ed) The population structure of vegetation. (W.) Junk, The Hague, pp 515–549Google Scholar
  22. Kleijn D, Treier UA, Müller-Schärer H (2005) The importance of nitrogen and carbohdrate storage for plant growth of the alpine herb Veratrum album. New Phytol 166:565–575PubMedCrossRefGoogle Scholar
  23. Körner C (2003) Alpine plant life, 2nd edn. Springer, Berlin Heidelberg New YorkGoogle Scholar
  24. de Kroon H, Huber H, Stuefer JF, van Groenendael JM (2005) A modular concept of phenotypic plasticity in plants. New Phytol 166:73–82PubMedCrossRefGoogle Scholar
  25. Lovett Doust J (1980) Experimental manipulation of patterns of resource allocation in the growth cycle and reproduction of Smyrnium olustratum L. Biol J Linn Soc 13:155–166CrossRefGoogle Scholar
  26. Meloche CG, Diggle PK (2003) The pattern of carbon allocation supporting growth of preformed shoot primordia in Acomastylis rossii (Rosaceae). Am J Bot 90:1313–1320CrossRefGoogle Scholar
  27. Meusel H, Jaeger E, Rauschert S, Weinert E (1978) Vergleichende Chorologie der zentraleuropaeischen Flora. Band II. Fischer, JenaGoogle Scholar
  28. Mössler JG (1833) Mösslers Handbuch der Gewächskunde. 1. Band 2. Abteilung 3. Auflage, JF Hammerich, AltonaGoogle Scholar
  29. Onipchenko V (ed) (2004) Alpine vegetation ecology of the western Caucasus. Kluwer, DordrechtGoogle Scholar
  30. Orshan G (1989) Plant pheno-morphological studies in Mediterranean type ecosystems. Kluwer, DordrechtGoogle Scholar
  31. Othen B, Wehrmeyer A (2004) Seasonal dynamics of non-structural carbohydrates in bulbs & shoots of the geophyte Galanthus nivalis. Physiol Plant 120:529–536CrossRefGoogle Scholar
  32. Packham JR, Harding DJL (1982) Ecology of woodland processes. Edward Arnold, LondonGoogle Scholar
  33. Pantis JD (1993) Biomass and nutrient allocation patterns in the Mediterranean geophyte Asphodelus aestivus Brot (Thessaly, Greece). Acta Oecol 14:489–500Google Scholar
  34. Pigliucci M (2001) Phenotypic plasticity: beyond nature and nurture. The Johns Hopkins University Press, BaltimoreGoogle Scholar
  35. Rauh M (1950) Morphologie der Nutzpflanzen. Quelle & Meyer, HeidelbergGoogle Scholar
  36. Rockwood LL, Lobstein MB (1994) The effects of experimental defoliation on reproduction in four species of herbaceous perennials from northern Virginia. Castanea 59:41–50Google Scholar
  37. Salisbury EJ (1925) The structure of woodlands. Veroff Geobot Inst Rübel Zürich 3:334–354Google Scholar
  38. Schlichting CD, Pigliucci M (1998) Phenotypic evolution. A reaction norm perspective. Sinauer Associates Inc., Sunderland Google Scholar
  39. Sohn JJ, Policansky D (1977) The costs of reproduction in the mayapple Podophyllum peltatum (Berberidaceae). Ecology 58:1366–1374CrossRefGoogle Scholar
  40. Suzuki JI, Stuefer JF (1999) On the ecological and evolutionary significance of storage in clonal plants. Plant Species Biol 14:11–17CrossRefGoogle Scholar
  41. von Willert DJ, Eller BM, Werger MJA, Brinckmann E, Ihlenfeldt HD (1992) Life strategies of succulents in deserts. Cambridge University Press, CambridgeGoogle Scholar
  42. Werger MJA, van Laar EMJM (1985) Seasonal changes in the structure of the herb layer of a deciduous woodland. Flora 176:351–364Google Scholar
  43. Whigham DF (1974) An ecological life-history study of Uvularia perfoliata. Am Midl Nat 91:343–359CrossRefGoogle Scholar
  44. Whigham DF (1990) The effects of experimental defoliation on the growth and reproduction of a woodland orchid, Tipularia discolor. Can J Bot 68:1812–1816Google Scholar
  45. Wijensinghe DK, Whigham DF (1997) Cost of producing clonal offspring and the effects of plant size on population dynamics of the woodland herb Uvularia perfoliata (Liliaceae). J Ecol 85:907–919CrossRefGoogle Scholar
  46. Willems JH, Bobbink R (1990) Spatial processes in the succession of chalk grassland on old fields in the Netherlands. In: Krahulec F, Agnew AD, Willems JH (eds) Spatial processes in plant communities. SPB Academic Publishing, The Hague, pp 237–249Google Scholar
  47. Worley AC, Harder LD (1999) Consequences of preformation for dynamic resource allocation by a carnivorous herb, Pinguicula vulgaris (Lentibulariaceae). Am J Bot 86:1136–1145PubMedCrossRefGoogle Scholar
  48. Wyka T (1999) Carbohydrate storage and use in an alpine population of the perennial herb, Oxytropis sericea. Oecologia 120:198–208CrossRefGoogle Scholar
  49. Wyka T (2000) Effects of nutrients on growth rate and carbohydrate storage in Oxytropis sericea: A test of the carbon accumulation hypothesis. Int J Plant Sci 161:381–386PubMedCrossRefGoogle Scholar
  50. Zimmerman JK, Whigham DF (1992) Ecological functions of carbohydrates stored in corms of Tipularia discolor (Orchidaceae). Funct Ecol 6:575–561CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2005

Authors and Affiliations

  1. 1.Department of Plant EcologyUtrecht UniversityUtrechtThe Netherlands
  2. 2.Department of EcologyRadboud University NijmegenNijmegenThe Netherlands

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