Microphylls and megaphylls

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In plant anatomy and evolution a microphyll (or lycophyll) is a type of plant leaf with one single, unbranched leaf vein. [1] Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concept of a microphyll, the leaf 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, have multiple veins within the leaf and leaf gaps above them in the stem.

Contents

Leaf vasculature

Microphylls contain a single vascular trace. Illustration Isoetes lacustris0.jpg
Microphylls contain a single vascular trace.

The clubmosses and horsetails have microphylls, as in all extant species there is only a single vascular trace in each leaf. [2] These leaves are narrow because the width of the blade is limited by the distance water can efficiently diffuse cell-to-cell from the central vascular strand to the margin of the leaf. [3] Despite their name, microphylls are not always small: those of Isoëtes can reach 25 centimetres in length, and the extinct Lepidodendron bore microphylls up to 78 cm long. [2]

Evolution

The enation theory of microphyll evolution posits that small outgrowths, or enations, developed from the side of early stems (such as those found in the Zosterophylls). [4] Outgrowths of the protostele (the central vasculature) later emerged towards the enations (as in Asteroxylon ), [4] and eventually continued to grow fully into the leaf to form the mid-vein (such as in Baragwanathia [4] ). [1] The fossil record appears to display these traits in this order, [4] but this may be a coincidence, as the record is incomplete. The telome theory proposes instead that both microphylls and megaphylls originated by the reduction; microphylls by reduction of a single telome branch, and megaphylls by evolution from branched portions of a telome. [4]

The simplistic evolutionary models, however, do not correspond well to evolutionary relationships. 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 had leaves with complex venation, and their current state is a result of secondary simplification. [5] Some gymnosperms bear needles with only one vein, but these evolved later from plants with complex leaves. [1]

An interesting case is that of Psilotum , which has a (simple) protostele, and enations devoid of vascular tissue. Some species of Psilotum have a single vascular trace that terminates at the base of the enations. [2] 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. [6]

It is not clear whether leaf gaps are a homologous trait of megaphyllous organisms or have evolved more than once. [1]

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.

See also

Related Research Articles

<span class="mw-page-title-main">Fern</span> Class of vascular plants

The ferns are a group of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the branched sporophyte is the dominant phase.

<span class="mw-page-title-main">Vascular plant</span> Phylum of plants with xylem and phloem

Vascular plants, or collectively the phylum Tracheophyta, form a large group of land plants that have lignified tissues for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue to conduct products of photosynthesis. Vascular plants include the clubmosses, horsetails, ferns, gymnosperms, and angiosperms. Scientific names for the group include Tracheophyta, Tracheobionta and Equisetopsida sensu lato. Some early land plants had less developed vascular tissue; the term eutracheophyte has been used for all other vascular plants, including all living ones.

<i>Selaginella</i> Genus of vascular plants in the family Selaginellaceae

Selaginella is the sole genus in the family Selaginellaceae, the spikemosses or lesser clubmosses, a kind of vascular plant.

<span class="mw-page-title-main">Lycophyte</span> Broadly circumscribed group of spore bearing plants

The lycophytes, when broadly circumscribed, are a group of vascular plants that include the clubmosses. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldest lineages of extant (living) vascular plants; the group contains extinct plants that have been dated from the Silurian. Lycophytes were some of the dominating plant species of the Carboniferous period, and included the tree-like Lepidodendrales, some of which grew over 40 metres (130 ft) in height, although extant lycophytes are relatively small plants.

<i>Psilotum</i> Genus of ferns in the family Psilotaceae

Psilotum is a genus of fern-like vascular plants. It is one of two genera in the family Psilotaceae commonly known as whisk ferns, the other being Tmesipteris. Plants in these two genera were once thought to be descended from the earliest surviving vascular plants, but more recent phylogenies place them as basal ferns, as a sister group to Ophioglossales. They lack true roots and leaves are very reduced, the stems being the organs containing photosynthetic and conducting tissue. There are only two species in Psilotum and a hybrid between the two. They differ from those in Tmesipteris in having stems with many branches and a synangium with three lobes rather than two.

<span class="mw-page-title-main">Osmundaceae</span> Family of ferns

Osmundaceae is a family of ferns containing four to six extant genera and 18–25 known species. It is the only living family of the order Osmundales in the class Polypodiopsida (ferns) or in some classifications the only order in the class Osmundopsida. This is an ancient and fairly isolated group that is often known as the "flowering ferns" because of the striking aspect of the ripe sporangia in Claytosmunda, Osmunda, Osmundastrum, and Plensium. In these genera the sporangia are borne naked on non-laminar pinnules, while Todea and Leptopteris bear sporangia naked on laminar pinnules. Ferns in this family are larger than most other ferns.

<span class="mw-page-title-main">Embryophyte</span> Subclade of green plants, also known as land plants

The Embryophyta, or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sister of the Zygnematophyceae. The Embryophyta consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and flowering plants. The land plants have diplobiontic life cycles and it is accepted now that they emerged from freshwater, multi-celled algae.

<span class="mw-page-title-main">Frond</span> Collection of leaflets on a plant

A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the large leaves of cycads, as well as palms (Arecaceae) and various other flowering plants, such as mimosa or sumac. "Frond" is commonly used to identify a large, compound leaf, but if the term is used botanically to refer to the leaves of ferns and algae it may be applied to smaller and undivided leaves.

In a vascular plant, the stele is the central part of the root or stem containing the tissues derived from the procambium. These include vascular tissue, in some cases ground tissue (pith) and a pericycle, which, if present, defines the outermost boundary of the stele. Outside the stele lies the endodermis, which is the innermost cell layer of the cortex.

<span class="mw-page-title-main">Equisetidae</span> Subclass of ferns

Equisetidae is one of the four subclasses of Polypodiopsida (ferns), a group of vascular plants with a fossil record going back to the Devonian. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.

<span class="mw-page-title-main">Sporophyll</span>

A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls bear either megasporangia and thus are called megasporophylls, or microsporangia and are called microsporophylls. The overlap of the prefixes and roots makes these terms a particularly confusing subset of botanical nomenclature.

<span class="mw-page-title-main">Psilotaceae</span> Family of ferns

Psilotaceae is a family of ferns consisting of two genera, Psilotum and Tmesipteris with about a dozen species. It is the only family in the order Psilotales.

<i>Asteroxylon</i> Extinct genus of spore-bearing plants

Asteroxylon is an extinct genus of vascular plants of the Division Lycopodiophyta known from anatomically preserved specimens described from the famous Early Devonian Rhynie chert and Windyfield chert in Aberdeenshire, Scotland. Asteroxylon is considered a basal member of the Lycopsida.

<span class="mw-page-title-main">Drepanophycales</span> Extinct order of spore-bearing plants

Drepanophycales is an order of extinct lycophyte plants of Late Silurian to Late Devonian age, found in North America, China, Russia, Europe, and Australia. Sometimes known as the Asteroxylales or Baragwanathiales.

<span class="mw-page-title-main">Evolutionary history of plants</span> History of plants

The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats of unicellular archaeplastids evolved through endosymbiosis, through multicellular marine and freshwater green algae, to spore-bearing terrestrial bryophytes, lycopods and ferns, and eventually to the complex seed-bearing gymnosperms and angiosperms of today. While many of the earliest groups continue to thrive, as exemplified by red and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones; for example, the ascendance of flowering plants over gymnosperms in terrestrial environments.

Enations are scaly leaflike structures, differing from leaves in their lack of vascular tissue. They are created by some leaf diseases and occur normally on Psilotum. Enations are also found on some early plants such as Rhynia, where they are hypothesized to have aided in photosynthesis.

<span class="mw-page-title-main">Polysporangiophyte</span> Spore-bearing plants with branched sporophytes

Polysporangiophytes, also called polysporangiates or formally Polysporangiophyta, are plants in which the spore-bearing generation (sporophyte) has branching stems (axes) that bear sporangia. The name literally means 'many sporangia plant'. The clade includes all land plants (embryophytes) except for the bryophytes whose sporophytes are normally unbranched, even if a few exceptional cases occur. While the definition is independent of the presence of vascular tissue, all living polysporangiophytes also have vascular tissue, i.e., are vascular plants or tracheophytes. Extinct polysporangiophytes are known that have no vascular tissue and so are not tracheophytes.

<span class="mw-page-title-main">Euphyllophyte</span> Clade of vascular plants

The euphyllophytes are a clade of plants within the tracheophytes. The group may be treated as an unranked clade, a division under the name Euphyllophyta or a subdivision under the name Euphyllophytina. The euphyllophytes are characterized by the possession of true leaves ("megaphylls"), and comprise one of two major lineages of extant vascular plants. As shown in the cladogram below, the euphyllophytes have a sister relationship to the lycopodiophytes or lycopsids. Unlike the lycopodiophytes, which consist of relatively few presently living or extant taxa, the euphyllophytes comprise the vast majority of vascular plant lineages that have evolved since both groups shared a common ancestor more than 400 million years ago. The euphyllophytes consist of two lineages, the spermatophytes or seed plants such as flowering plants (angiosperms) and gymnosperms, and the Polypodiophytes or ferns, as well as a number of extinct fossil groups.

<span class="mw-page-title-main">Leaf</span> Photosynthetic part of a vascular plant

A leaf is a principal appendage of the stem of a vascular plant, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, flower, and fruit collectively form the shoot system. In most leaves, the primary photosynthetic tissue is the palisade mesophyll and is located on the upper side of the blade or lamina of the leaf but in some species, including the mature foliage of Eucalyptus, palisade mesophyll is present on both sides and the leaves are said to be isobilateral. Most leaves are flattened and have distinct upper (adaxial) and lower (abaxial) surfaces that differ in color, hairiness, the number of stomata, the amount and structure of epicuticular wax and other features. Leaves are mostly green in color due to the presence of a compound called chlorophyll which is essential for photosynthesis as it absorbs light energy from the sun. A leaf with lighter-colored or white patches or edges is called a variegated leaf.

The barinophytes are a group of extinct vascular plants (tracheophytes). Their relationship with other vascular plants is unclear. They have been treated as the separate class Barinophytopsida, the order Barinophytales of uncertain class and as a family or clade Barinophytaceae within the zosterophylls. They have also been considered to be possible lycopodiopsids.

References

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