title: Impact of spatial constraints during seed germination on the evolution of angiosperm cotyledons: a case study from tropical Hydatellaceae (Nymphaeales).
abstract: OBJECTIVE
  A bipolar embryo with cotyledons is a characteristic feature that appeared early in the evolution of seed plants. Cotyledon number is an important character in angiosperm classification. We explore the links between functional aspects of seed germination and the number and location of the cotyledons, using as a model the early-divergent angiosperm family Hydatellaceae, in which seedlings are superficially monocot-like. •
  
  
  METHODS
  Seedlings of two species of tropical Hydatellaceae were studied using light and scanning electron microscopy. •
  
  
  RESULTS
  Seedlings of Trithuria cowieana bear two free cotyledons. Each cotyledon possesses a green, filiform, vascularized blade that resembles subsequent leaves, and a basal, nonvascularized, haustorial outgrowth that remains in close contact with the endosperm. Seedlings of Trithuria konkanensis have two free cotyledonary haustoria inserted close to each other and a leaf blade probably belonging to one of the cotyledons. The cotyledonary node elongates between the haustoria and the leaf blade to form a mesocotyl. •
  
  
  CONCLUSIONS
  To date, the absence or presence of a cotyledonary tube represents the only known qualitative morphological difference between the two major clades of Hydatellaceae. Cotyledons with a haustorium and leaf blade are unusual at the scale of seed plants and probably evolved due to homeosis. The mesocotyl of T. konkanensis resembles that of grasses and sedges. Seedling diversity in Hydatellaceae and other seed plants is linked with the principal physical and spatial constraint of their embryo structure, with the primary root and shoot apical meristems located at opposite poles, and haustorial cotyledon tips.
authors.affiliation: Department of Higher Plants, Faculty of Biology, Lomonosov Moscow State University, Moscow 119234, Russia. sokoloff-v@yandex.ru
authors.forename: Dmitry D
authors.initials: DD
authors.lastname: Sokoloff
authors.forename: Margarita V
authors.initials: MV
authors.lastname: Remizowa
authors.forename: Anton S
authors.initials: AS
authors.lastname: Beer
authors.forename: Shrirang R
authors.initials: SR
authors.lastname: Yadav
authors.forename: Terry D
authors.initials: TD
authors.lastname: Macfarlane
authors.forename: Margaret M
authors.initials: MM
authors.lastname: Ramsay
authors.forename: Paula J
authors.initials: PJ
authors.lastname: Rudall
mesh_terms: D005075:Biological Evolution; D018548:Cotyledon / Q000235:genetics / Q000502:physiology*; D017753:Ecosystem; D018525:Germination; D019684:Magnoliopsida / Q000235:genetics / Q000502:physiology* / Q000648:ultrastructure; D036226:Seedlings / Q000648:ultrastructure; D012639:Seeds; D014329:Tropical Climate
keywords: Trithuria; embryo; heterocotyly; homeosis; homology; mesocotyl; monocotyledons; phyllotaxy; seed plants; syncotyly