Liverwort

Liverworts
Temporal range: 472–0 Ma Mid-Ordovician to present
Metzgeria furcata, a species with a flat thallus
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Embryophytes
Clade: Setaphyta
Division: Marchantiophyta
Stotler & Crand.-Stotl., 1977 emend. 2000
Type genus
Marchantia L., 1753
Classes and orders
Synonyms
  • Hepaticae Juss., 1789
  • Marchantiophytina Doweld, 2001
  • Hepaticophyta Crand.-Stotl et Stotler, 2000
  • Hepatophyta Stotler et Crand.-Stotl, 1977
  • Jungermanniophyta

Liverworts are non-vascular land plants in the division Marchantiophyta (/mɑːrˌkæn.ti.ˈɒf.ə.tə, -oʊ.ˈfaɪ.tə/ ⓘ). Liverworts, mosses, and hornworts are collectively known as bryophytes. Like mosses and hornworts, liverworts have a gametophyte-dominant life cycle in which the sporophyte is dependent on the gametophyte.

Some liverworts have a flat branching thallus, while others are leafy. They are distinguished from mosses by their single-celled rhizoids, the absence of a midrib or costa in the leaves, and the presence of membrane-bound complex oil bodies.

There are an estimated 9,000 species of liverworts, most of them leafy. They occur in many terrestrial ecosystems worldwide, especially in moist tropical regions, although some are adapted to deserts and other dry environments.

Physical characteristics

Description

Most liverworts are small, from 2 to 20 millimetres (0.08 to 0.8 in) wide with individual plants less than 10 centimetres (4 in) long, so they are often overlooked. The most familiar liverworts, in the order Marchantiales, consist of a prostrate, flattened, ribbon-like or branching structure called a thallus (plant body). However, most liverworts produce flattened stems with overlapping scales or leaves in two or more ranks, the middle rank is often conspicuously different from the outer ranks; these leafy liverworts are in the order Jungermanniales.

Recognition

Liverworts can most reliably be distinguished from similar-looking mosses by their single-celled rhizoids. Other differences are not universal for all mosses and liverworts, but the lack of clearly differentiated stem and leaves in species with a thallus, or deeply lobed or segmented leaves and the presence of leaves arranged in three ranks in leafy species, the absence of a midrib or costa in the leaves, and frequent dichotomous branching all point to the plant's being a liverwort. With a few exceptions, all liverworts undergo polyplastidic meiosis, in contrast to mosses and hornworts which have monoplastidic meiosis. Unlike any other embryophytes, most liverworts contain complex oil bodies containing isoprenoids and surrounded by membranes in some of their cells.

Life cycle

Sexual

Liverworts, like other bryophytes, have a gametophyte-dominant life cycle, with the sporophyte dependent on the gametophyte. The sporophytes of many but not all liverworts are non-photosynthetic. Cells of the gametophyte plant each contain only a single set of genetic information, so liverworts are haploid for the majority of their life cycle. This contrasts sharply with the pattern in seed plants, where the haploid generation is represented only by the tiny pollen and ovule, while the diploid generation is the familiar plant. Another unusual feature is that the sporophytes (i.e. the diploid body) are very short-lived, withering away soon after releasing their spores.

The life of a liverwort starts when a haploid spore germinates to produce a transitory protonema, which is either a mass of thread-like filaments or a flattened thallus. The protonema grows into a mature gametophyte plant that produces the sex organs. The male organ is known as an antheridium; it produces sperm cells. The female organ is known as an archegonium. Each archegonium has a slender hollow tube or neck down which the sperm swim to reach the egg cell.

Liverwort species may be either dioicous or monoicous. In dioicous liverworts, female and male sex organs are borne on different and separate gametophyte plants. In monoicous liverworts, the two kinds of reproductive structures are borne on different branches of the same plant. In either case, the sperm must move from the antheridia where they are produced to the archegonium where the eggs are held. The sperm of liverworts have two tail-like flagella that enable them to swim short distances.

When sperm reach the archegonia, fertilisation occurs, leading to the production of a diploid sporophyte. After fertilisation, the immature sporophyte within the archegonium develops three distinct regions: (1) a foot, which anchors the sporophyte in place and receives nutrients from its "mother" plant, (2) a spherical or ellipsoidal capsule, inside which the spores will be produced for dispersing to new locations, and (3) a seta (stalk) which connects the capsule to the foot. The mature capsule splits, scattering the spores.

Asexual

Most liverworts are capable of asexual reproduction, where it is common and indeed the usual mode of reproduction. For example, in Riccia, when the older parts of the forked thalli die, the younger tips become separate individuals.

Some thallose liverworts such as Marchantia polymorpha and Lunularia cruciata produce small disc-shaped gemmae in shallow cups. Marchantia gemmae can be dispersed up to 120cm by rain splashing into the cups. In Metzgeria, gemmae grow at thallus margins. M. polymorpha is a common weed in greenhouses, often covering the entire surface of containers, as it spreads rapidly by the dispersal of its gemmae.

Ecology

Liverworts grow in many ecosystems across the planet except the sea and excessively dry environments, or those exposed to high levels of direct solar radiation. They are most common (both in numbers and species) in moist tropical areas.

Thalloid liverworts host symbiotic glomeromycete fungi. Species in the Aneuraceae, however, associate with basidiomycete fungi of the genus Tulasnella, while leafy liverworts host symbiotic basidiomycete fungi of the genus Serendipita.

Evolution

Epiphytic thalloid liverworts evolved during the Triassic to the Cretaceous.

External phylogeny

Traditionally, the liverworts were grouped together with the mosses and hornworts in the Division Bryophyta, within which the liverworts made up the class Hepaticae (also called Marchantiopsida). Somewhat more recently, the liverworts were given their own division (Marchantiophyta), as bryophytes became considered to be paraphyletic. However, phylogenetic evidence indicates that liverworts are likely part of a clade ("Bryophyta sensu lato" or "Bryophyta Schimp.") alongside mosses and hornworts. Hence, it has been suggested that the liverworts should be de-ranked to a class called Marchantiopsida. In addition, there is strong phylogenetic evidence that liverworts and mosses form a subclade, Setaphyta.

Two of the most likely models for bryophyte evolution
'Monophyletic bryophytes' 'Liverworts plus mosses – basal'
embryophytes
embryophytes

Fossil record

Radula cretacea, a liverwort fossil in Cretaceous amber from Burma

Stomata appear to be ancestral, and to have been lost in the liverwort lineage. Among the earliest fossils believed to be liverworts are compression fossils of Pallaviciniites from the Upper Devonian of New York. These fossils resemble modern species in the Metzgeriales. Another Devonian fossil, Protosalvinia, looks like a liverwort, but its relationship to other plants is uncertain, so it may not belong to the Marchantiophyta. In 2007, the oldest fossil species assignable to the liverworts was Metzgeriothallus sharonae from the Givetian (Middle Devonian) of New York. In 2010, five different types of fossilized liverwort spores were found in Argentina, dating to the much earlier Middle Ordovician, around 470 million years ago. In 2012, the earliest simple thalloid liverwort fossil, Riccardiothallus devonicus, was found in the Devonian of Yunnan, China. Total-evidence dating in 2023 combining morphological analysis of eight fossil taxa and genomic analysis of 56 extant species suggests that the Marchantiopsida radiated in the Middle Ordovician, while the Ricciaceae radiated in the Middle Jurassic. The most recent common ancestor of the complex thalloid liverworts was reconstructed as resembling Blasia with a flat thallus and a sporophyte capsule that split into four.

Internal phylogeny

Bryologists classify liverworts in the division Marchantiophyta. This divisional name is based on the name of the genus Marchantia. In addition, the liverworts have sometimes been called Hepaticophyta, from Greek ‘ἡπατικός "hepatikos", of the liver.

The Marchantiophyta may be subdivided into three classes:

Forrest 2006 Cole, Hilger & Goffinet 2021

A classification by Söderström et al. 2016

There are about 9000 species of liverworts, at least 85% of which belong to the leafy group. The first liverwort genome sequence, from Marchantia polymorpha, was reported in 2017 although a few genes had already been identified and characterized.

Relationship with humans

"Hepaticae" from Ernst Haeckel's Kunstformen der Natur, 1904

In ancient times, it was believed that liverworts cured diseases of the liver, hence their name, which in its Old English form liferwyrt means 'liver plant'. The archaic relationship of plant form to function is based on the medieval doctrine of signatures. At the start of the 20th century, the German zoologist Ernst Haeckel depicted liverworts in an elaborately symmetrical colour plate of the "Hepaticae" in his book Kunstformen der Natur.

Riccia fluitans (floating crystalwort) is an aquatic thallose liverwort sold for use in aquariums. Its slender branches float on the water's surface and provide habitat for both small invertebrates and the fish that feed on them.

Bryophyte genomics offers economic possibilities. Liverworts can be grown in petri dishes. Their gemmae provide large quantities of ready-made asexual clones. There is an efficient genetic transformation system; transgenic Marchantia polymorpha lines can be generated in 4 weeks. Many genetic engineering tools designed for angiosperms work in liverworts, including CRISPR systems. M. polymorpha could form a rapid system for testing plant genetic engineering and synthetic biology before moving into angiosperms. It may be practical to grow it in hydroponic systems and harvest high-value recombinant proteins. M. polymorpha had by 2026 become the model organism for molecular studies of liverworts.

References

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