Nonvascular Plants: Mosses, Liverworts, & Hornworts

Characteristics of nonvascular plants

embryophytes with no vascular tissue

multicellular sporangia and gametangia

reproductive cells surrounded by one or several layers of sterile cells

bodies composed of true parenchyma derived from apical meristems

terrestrial and have a cuticle over much of their bodies and many have stomata

life cycle

alternation of heteromorphic generations

sporophyte

gametophyte

in flowering plants they are large with leaf and roots

tiny and occur inside pollen grains and ovules

however they are the larger more prominent generation in nonvascular

in nonvascular they are smaller, more temporary and very inconspicuous

green moss

absorb minerals from gametophytes and remain attached to them

never independent of of the gametophyte, never branch, and never have leaves

visible if you examine moss closely, and look like brown or green hairs

wont grow to be very large, but being small provides certain advantages

selective advantage in habitats

Classification of nonvascular plants

not known how closely related mosses, liverworts, and hornworts are yet

they have features in common, but also differ in certain aspects

treated as three distinct divisions

mosses= Bryophyta

liverworts= Hepatophyta

hornworts= Anthocerotophyta

Division of Bryophya: mosses

Gametophyte generation

mosses= ubiquitous, occurring in all parts of the world and in almost every environment

Morphology

leafy stems = gametophores which grow closely together, tightly appressed, and form dense mounds

cool wet environments produces plants more open and loose

all moss stems have leaves

did not evolve from same structures as vascular plants stems and leaves did

gametophores grow from an apical meristem that contains a apical cell

tissues and leaves in precise arrangements

cuticle occurs only on top part of most moss leaves making the underside capable of absorbing water directly

lack of cuticle= no protection against desiccation

no stomata on moss leaves

moss stems are always slender and have little tissue differentiation

surface not to different from underlying so no epidermis

stem tissues all called cortex, may be uniform in all parts

outer cells more narrow with thickened walls

inner cells are larger and more parenchymatous and chlorophyllous

Water Transport

in some mosses, primarily the family Polytrichaceae, the inner most cortex has cells composed of hydroids

conduct water and dissolved minerals

elongated cells that lose cytoplasm when mature and end walls are partially digested away

aligned with those above and below

species with hydroids usually have leptoids then

resemble sieve cells

no nuclei, prominent gap junctions between cells, elongated and retain nuclei

majority of mosses lack hydroids and leptoids

water conducted along the exterior of stems via capillary action

leaves and stems so small they form spaces narrow enough to act as capillary channels

at the base of the stem is rhizoids

no leptoids means water transported between parenchyma cells

small, multicellular thrichome like structures

anchor stems by penetrating substrate

Development

gametophore growth begins when a spore germinates and sends out long slender, chlorophyllous cell

cell undergoes mitosis and produces a branched system= protonema

Reproduction

gametophore produces gametangia

all mosses are oogamous= every species has small bifilagellate sperm cells and large nonmotile egg cells

sperm produced in microgametangia called antheridia

eggs occur in megagametangia called archegonia

both occur on same gametophore in bisexual species

other species have male and female gametophores

special mechanisms used to get sperm to the egg

Sporophyte generation

moss gametophytes are both large and photosynthetic and they support the sporophyte its entire life

zygote undergoes a transverse division and develops into a small bulbous tissue called the foot

foot of gametophore used to absorb sugars, minerals, and water

cells of foot are transfer cells in m any species

upper cells grow by cell division and become simple apical sporangium= capsule

between the foot and the stalk is the seta

sporophyte is structurally simple

true epidermis with stomata

apex of sporangium is a cap like lid called operculum

breakage of cell from operculum separating from sporangium= peristome teeth

teeth trap spores when humid, and opening to release when air is dry

sporangium covered by calyptra

Metabolism and ecology

mechanisms used to retain water due to inability of mosses to retain water

many species grow in permanently moist environments

some mosses tolerant of desiccation and drying out does not damage them

as long as 30% of their weight is water, then they remain dormant but alive

dessicated mosses resistant to high or low temps and UV light

grow on hard or impervious surfaces

important in later establishment of other species

Division of Hepatophyta: Liverworts

small plants that have alternation of heteromorphic generations

few species ever become large

some species are leafy and greatly resemble mosses

others form small, solid, ribbon like gametophytes with a less conspicuous sporophyte then mosses

Gametophyte generation

divided into two basic groups

leafy liverworts

thallose liverworts

in both the gametophyte generation initiated when spores germinate and establish a protonematal phase

gametophore resembles that of moss, slender stem with leaves

leaves differ from moss though however

show less resemblance to mosses

not leafy at all but rather flat and ribbon like or heart shaped and bilaterally symmetrical

body referred as a thallus which has no roots, stems, or leaves

much thicker then leafy

many cells contain oil droplets next to substrate and those on the side away from substrate have more chlorophyll

cells arranged in a aerenchyma with large air chambers that open to exterior via large air pores

not stomata

both may either be bisexual, or unisexual

Sporophyte generation

little variability and basic morphology is like that of mosses

differ in fundamental structure though

liverwort lack a columella

dehiscence occurs via 4 longitudinal slits, not by an operculum

no peristome teeth

cells in sporangium do not undergo meiosis, rather they differentiate into elaters

Division Anthocerotophyta: Hornworts

small, inconspicuous thalloid plants that grow on moist soil, hidden by grasses and other herbs

resemble thalloid liverworts

alternation of heteromorphic generations occurs

large, single chloroplast per cell

chloroplasts have a pyrenoid

Gametophyte generation

gametophores are very thin

gametophyte is parenchymous and does not tolerate drying

internally gametophytes have numerous chambers

chambers start off as mucilage chambers, but become invaded by bacteria to form a symbiotic relationship

special mucilage chamber used to develop antheridia

archegonia formed via superficial cells

Sporophyte generation

similarities hard to find between mosses and liverworts

hornworts have a foot embedded in gametophore tissue, but no seta or discrete sporangium

sporangium is a long, horn like cylinder

some species have stomata

spores are green, yellow, brown, or black

have a columella as in mosses, and have elaters like liverworts, however these are multicellular unlike liverworts

nonvascular plants

mosses

liverworts

liverwort life cycle

hornworts

hornworts 2