Microphyll

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This article is about leaf vasculature and evolution. For other uses, see Microphyll (disambiguation).
Megaphyll leaves display a complex network of veins.

Microphyll is a type of plant leaf, which has traditionally been defined as "an appendage supplied by a single, unbranched vein".[1] In the classical concept of a microphyll, this vein emerges from the protostele without leaving a leaf gap. Leaf gaps are small areas above the node of some leaves where there is no vascular tissue, as it has all been diverted to the leaf. Megaphylls, in contrast, are characterised by multiple veins. By this definition, the whisk ferns, clubmosses and horsetails have microphylls, as in all extant species there is only a single vascular trace in each leaf.[2] Despite their name, microphylls are not always small: those of Isoëtes can reach 20–100 centimetres in length, and the extinct Lepidodendron bore microphylls over a metre long.[citation needed]

Evolution of leaves: Microphylls and Megaphylls[edit]

Microphylls contain a single vascular trace.

The "Enation theory" of microphyll development posits that small outgrowth, or enations, developed from the side of early axes (such as those found in the Zosterophylls).[3] Outgrowths of the protostele later emerged towards the enations (as in Asteroxylon),[3] and eventually continued to grow fully into the leaf to form the mid-vein (such as in Baragwanathia[3]).[1] The fossil record appears to display these traits in this order,[3] but this may be a coincidence, as the record is very incomplete: microphylls may have originated by the reduction of a telome branch.[3]

Exceptions[edit]

The simplistic models described above do not apply to all organisms. Some genera of ferns display complex leaves that are attached to the pseudostele by an outgrowth of the vascular bundle, leaving no leaf gap.[1] Horsetails (Equisetum) bear only a single vein, and appear to be microphyllous; however, the fossil record suggests that their forebears bore leaves with complex venation, and the current state is a result of secondary simplification.[4] Some gymnosperms bear needles with only one vein, but these evolved later from plants with complex leaves.[1]

Psilotum has secondarily lost leaves, and bears enations resembling those of early land plants.

An interesting case is that of Psilotum, which bears a protostele, and enations devoid of vascular tissue. Consequently, Psilotum was long thought to be a "living fossil" closely related to early land plants (Rhyniophytes). However, genetic analysis has shown Psilotum to be a reduced fern.[5]

To make matters worse, there is also some debate about whether leaf gaps are unique and/or common to megaphyllous organisms.[1]

This ambiguity leaves it difficult to distinguish between two competing hypotheses: that microphylls evolved via the reduction of megaphylls, and that they evolved independently, from enations. While the simple definitions (microphylls: one vein, macrophylls: more than one) can still be used in modern botany, the evolutionary history is harder to decipher.

References[edit]

  1. ^ a b c d e Kaplan, D.R. (2001). "The Science of Plant Morphology: Definition, History, and Role in Modern Biology". American Journal of Botany 88 (10): 1711–1741. doi:10.2307/3558347. JSTOR 3558347. PMID 21669604. 
  2. ^ Gifford E.M. & Foster, A.S. (1989). Morphology and evolution of vascular plants. WH Freeman, New York, USA. 
  3. ^ a b c d e WN Stewart & GW Rothwell (1993) Palaeobotany and the evolution of plants. 2nd edition. Cambridge University Press.
  4. ^ Taylor, T.N.; Taylor, E.L. (1993). The biology and evolution of fossil plants. 
  5. ^ Qiu, Y.L.; Palmer, J.D. (1999). "Phylogeny of early land plants: insights from genes and genomes". Trends in Plant Science 4 (1): 26–30. doi:10.1016/S1360-1385(98)01361-2. PMID 10234267.