Characteristics unique to Equisetum species among living vascular plants



1.  Lateral branch origin:

    Lateral branches of Equisetum stems arise between leaves (i.e. branches alternate with leaves).  In other plants with lateral (as distinguished from terminal) branching, branches originate in leaf axils (i.e. in the vertex of the upper angle between a leaf and the stem from which it arises)


2.  Stomatal structure:

    Dayanandan (1977) observed that Equisetum species "possess perhaps the most structurally complex stomata in the entire plant kingdom" (p. 175).  The stomata of equisetum are so unique that "a single well-preserved stomatal apparatus is all that is needed to identify the genus Equisetum (even the two subgenera) from among all other living plants" (Dayanandan, 1977).  The uniqueness of Equisetum stomata is the result of two characteristics (Dayanandan, 1977):  

1.) The two subsidiary cells overlie the guard cells completely, whereas in other plants the guard cells are the superficial cells.
2.) "The inner tangential wall of each subsidiary cell develops 7 to 24 ridge-like thickenings, a feature not found in any other genus." (Dayanandan, 1977)

These unique features are nicely shown in illustrations from the plates accompanying Milde (1867) of:
1.  a stoma of E. giganteum (subgenus Hippochaete)
2.  a stoma of E. bogotense (subgenus Equisetum)
note:  The subsidiary cells are illustrated transparently so that the kidney shaped guard cells can be seen beneath.

3.  Elaters on spores:

    Each Equisetum spore has four strap-like structures called elaters attached to the spore surface at a common point.  These elaters are hygroscopic (i.e. they expand and contract with changes in humidity) and probably function to help disperse the spores (Hauke, 1963).


4.  Silicon requirement:

    Among terrestrial plants, only the horsetails have been definitively shown to require Silicon as an essential, not simply beneficial, mineral nutrient (Epstein, 1999).  The requirement for silicon has been shown for Equisetum arvense ( Chen and Lewin, 1969 ) and for E. hyemale ( Hoffman and Hillson, 1979 ), so this requirement appears to hold for members of both subgenera within Equisetum .


5.  Rhizome architecture (?):

    Golub and Whetmore (1948) excavated the rhizome system of a colony of Equisetum arvense to a depth of 2 m and found five successive horizontal layers of rhizomes connected by vertical rhizomes.  This rhizome system extended below 2 m, but the investigators did not excavate further.  I suspect that this "tiered" rhizome architecture may be unique in the plant kingdom.  Indeed, other rhizomatous plants generally have but a single horizontal rhizome system layer ( Bell and Tomlinson, 1980 ).


If you have any comments or questions, please contact the author, Chad Husby ( chad.husby@fiu.edu or husby.1@osu.edu )

© Chad E. Husby 2002

Last updated October 21, 2002

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