Seedless Vascular Plants

Ferns and Other Seedless Vascular Plants

By the late Devonian period, plants had evolved vascular tissue, well-defined leaves, and root systems. With these advantages, plants increased in height and size. During the Carboniferous period, swamp forests of club mosses and horsetails—some specimens reaching heights of more than 30 m (100 ft)—covered most of the land. These forests gave rise to the extensive coal deposits that gave the Carboniferous its name. In seedless vascular plants, the sporophyte became the dominant phase of the lifecycle.

Water is still required for fertilization of seedless vascular plants, and most favor a moist environment. Modern-day seedless tracheophytes include club mosses, horsetails, ferns, and whisk ferns.

Phylum Lycopodiophyta: Club Mosses

The club mosses, or phylum Lycopodiophyta, are the earliest group of seedless vascular plants. They dominated the landscape of the Carboniferous, growing into tall trees and forming large swamp forests. Today’s club mosses are diminutive, evergreen plants consisting of a stem (which may be branched) and microphylls (Figure). The phylum Lycopodiophyta consists of close to 1,200 species, including the quillworts (Isoetales), the club mosses (Lycopodiales), and spike mosses (Selaginellales), none of which are true mosses or bryophytes.

Lycophytes follow the pattern of alternation of generations seen in the bryophytes, except that the sporophyte is the major stage of the lifecycle. The gametophytes do not depend on the sporophyte for nutrients. Some gametophytes develop underground and form mycorrhizal associations with fungi. In club mosses, the sporophyte gives rise to sporophylls arranged in strobili, cone-like structures that give the class its name. Lycophytes can be homosporous or heterosporous.

 In the photo, seed-like strobili are arranged around the slender stalks of a club moss.
In the club mosses such as Lycopodium clavatum, sporangia are arranged in clusters called strobili. (credit: Cory Zanker)

Phylum Monilophyta: Class Equisetopsida (Horsetails)

Horsetails, whisk ferns and ferns belong to the phylum Monilophyta, with horsetails placed in the Class Equisetopsida. The single genus Equisetum is the survivor of a large group of plants, known as Arthrophyta, which produced large trees and entire swamp forests in the Carboniferous. The plants are usually found in damp environments and marshes (Figure).

 In the photo, bushy horsetail plants grow in water.
Horsetails thrive in a marsh. (credit: Myriam Feldman)

The stem of a horsetail is characterized by the presence of joints or nodes, hence the name Arthrophyta (arthro- = "joint"; -phyta = "plant"). Leaves and branches come out as whorls from the evenly spaced joints. The needle-shaped leaves do not contribute greatly to photosynthesis, the majority of which takes place in the green stem (Figure).

 Photo shows a horsetail plant, which resembles a scrub brush, with a thick stem and whorls of thin leaves branching from the stem.
Thin leaves originating at the joints are noticeable on the horsetail plant. Horsetails were once used as scrubbing brushes and were nicknamed scouring brushes. (credit: Myriam Feldman)

Silica collects in the epidermal cells, contributing to the stiffness of horsetail plants. Underground stems known as rhizomes anchor the plants to the ground. Modern-day horsetails are homosporous and produce bisexual gametophytes.

Phylum Monilophyta: Class Psilotopsida (Whisk Ferns)

While most ferns form large leaves and branching roots, the whisk ferns, Class Psilotopsida, lack both roots and leaves, probably lost by reduction. Photosynthesis takes place in their green stems, and small yellow knobs form at the tip of the branch stem and contain the sporangia. Whisk ferns were considered an early pterophytes. However, recent comparative DNA analysis suggests that this group may have lost both vascular tissue and roots through evolution, and is more closely related to ferns.

 Photo shows a whisk fern with many green stems that have small knobs along their length.
The whisk fern Psilotum nudum has conspicuous green stems with knob-shaped sporangia. (credit: Forest & Kim Starr)

Phylum Monilophyta: Class Psilotopsida (Ferns)

With their large fronds, ferns are the most readily recognizable seedless vascular plants. They are considered the most advanced seedless vascular plants and display characteristics commonly observed in seed plants. More than 20,000 species of ferns live in environments ranging from tropics to temperate forests. Although some species survive in dry environments, most ferns are restricted to moist, shaded places. Ferns made their appearance in the fossil record during the Devonian period and expanded during the Carboniferous.

The dominant stage of the lifecycle of a fern is the sporophyte, which consists of large compound leaves called fronds. Fronds fulfill a double role; they are photosynthetic organs that also carry reproductive organs. The stem may be buried underground as a rhizome, from which adventitious roots grow to absorb water and nutrients from the soil; or, they may grow above ground as a trunk in tree ferns (Figure). Adventitious organs are those that grow in unusual places, such as roots growing from the side of a stem.

 Photo shows a potted tree fern.
Some specimens of this short tree-fern species can grow very tall. (credit: Adrian Pingstone)

The tip of a developing fern frond is rolled into a crozier, or fiddlehead (Figurea and Figureb). Fiddleheads unroll as the frond develops.

Fiddleheads at the top of a maturing fern curl into a structure that resembles their namesake.
Croziers, or fiddleheads, are the tips of fern fronds. (credit a: modification of work by Cory Zanker; credit b: modification of work by Myriam Feldman)

The lifecycle of a fern is depicted in Figure.

Art Connection

 The fern life cycle begins with a diploid (2n) sporophyte, which is the fern plant. Sporangia are round bumps that occur on the bottom of the leaves. Sporangia undergo mitosis to form haploid (1n) spores. The spores germinate and grow into a green gametophyte 1n that resembles lettuce. The gametophyte contains antheridia that produce, sperm and archegonia that produce eggs. Inside the archegonium the sperm fertilizes the egg, forming a diploid (2n) zygote. The zygote undergoes mitosis to form a 2n sporophyte, ending the cycle.
This life cycle of a fern shows alternation of generations with a dominant sporophyte stage. (credit "fern": modification of work by Cory Zanker; credit "gametophyte": modification of work by "Vlmastra"/Wikimedia Commons)

Which of the following statements about the fern life cycle is false?

  1. Sporangia produce haploid spores.
  2. The sporophyte grows from a gametophyte.
  3. The sporophyte is diploid and the gametophyte is haploid.
  4. Sporangia form on the underside of the gametophyte.

Link to Learning

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To see an animation of the lifecycle of a fern and to test your knowledge, go to the website.

Most ferns produce the same type of spores and are therefore homosporous. The diploid sporophyte is the most conspicuous stage of the lifecycle. On the underside of its mature fronds, sori (singular, sorus) form as small clusters where sporangia develop (Figure).

 The photo shows small bumps called sori on the underside of a fern frond.
Sori appear as small bumps on the underside of a fern frond. (credit: Myriam Feldman)

Inside the sori, spores are produced by meiosis and released into the air. Those that land on a suitable substrate germinate and form a heart-shaped gametophyte, which is attached to the ground by thin filamentous rhizoids (Figure).

 The photo shows a young sporophyte with a fan-shaped leaf growing from a lettuce-like gametophyte.
Shown here are a young sporophyte (upper part of image) and a heart-shaped gametophyte (bottom part of image). (credit: modification of work by "Vlmastra"/Wikimedia Commons)

The inconspicuous gametophyte harbors both sex gametangia. Flagellated sperm released from the antheridium swim on a wet surface to the archegonium, where the egg is fertilized. The newly formed zygote grows into a sporophyte that emerges from the gametophyte and grows by mitosis into the next generation sporophyte.

Career Connection

Landscape DesignerLooking at the well-laid parterres of flowers and fountains in the grounds of royal castles and historic houses of Europe, it’s clear that the gardens’ creators knew about more than art and design. They were also familiar with the biology of the plants they chose. Landscape design also has strong roots in the United States’ tradition. A prime example of early American classical design is Monticello: Thomas Jefferson’s private estate. Among his many interests, Jefferson maintained a strong passion for botany. Landscape layout can encompass a small private space, like a backyard garden; public gathering places, like Central Park in New York City; or an entire city plan, like Pierre L’Enfant’s design for Washington, DC.

A landscape designer will plan traditional public spaces—such as botanical gardens, parks, college campuses, gardens, and larger developments—as well as natural areas and private gardens. The restoration of natural places encroached on by human intervention, such as wetlands, also requires the expertise of a landscape designer.

With such an array of necessary skills, a landscape designer’s education includes a solid background in botany, soil science, plant pathology, entomology, and horticulture. Coursework in architecture and design software is also required for the completion of the degree. The successful design of a landscape rests on an extensive knowledge of plant growth requirements, such as light and shade, moisture levels, compatibility of different species, and susceptibility to pathogens and pests. Mosses and ferns will thrive in a shaded area, where fountains provide moisture; cacti, on the other hand, would not fare well in that environment. The future growth of individual plants must be taken into account, to avoid crowding and competition for light and nutrients. The appearance of the space over time is also of concern. Shapes, colors, and biology must be balanced for a well-maintained and sustainable green space. Art, architecture, and biology blend in a beautifully designed and implemented landscape.

 Photo shows a landscaped garden with a variety of flowers and bushes.
This landscaped border at a college campus was designed by students in the horticulture and landscaping department of the college. (credit: Myriam Feldman)
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