Charophyceae

Discipline Courses-I
Semester-I
Paper: Phycology and Microbiology
Unit-VIII
Lesson: Charophyceae
Lesson Developer: Geeta Sharma
College/Department: Shivaji College, University of Delhi

Institute of Lifelong Learning, University of Delhi

 Charophyceae

Table of Contents

Chapter: Charophyceae

 Introduction
 Chara sp.
 Habit
 Habitat
 Vegetative thallus
 Rhizoids
 Stem
 Cell structure
 Reproduction
 Asexual (vegetative)
 Sexual Reproduction
 Parthenogenesis
 Economic Importance
 Systematic Position and Affinities.

 Summary
 Exercise
 References
 Suggested Reading

Institute of Lifelong Learning, University of Delhi 1

 Charophyceae

Introduction

Members of Charophyceae are very closely related to the ancestory of land plants and
exhibit the useful information about the evolutionary process and the pattern of extinction.
It is an ancient group (known since Silurian period), that show the basic similarity in
pigmentation, metabolism, cell structure, reserve food, type of reproduction and occurrence
in fresh water habitat with green algae. There are six living genera and the remaining as
fossils. Charophyceae includes the genus Chara and Nitella commonly found in fresh water
bodies rich in minerals as epipelics. In this lesson the genus Chara is dealt in detail.

CHARA (Common name: Aquatic Horse Tail/ a mountain stream/ a stonewort )

Habit

Chara a fresh water, macroscopic, filamentous, multicellular benthonic epipelic alga.

Figure : Chara fragilis – W.M. of a plant collected from submerged condition.

Source: http://bobklips.com/bobs_website/CHARVULK-22Jul08-SPBI-PLE.jpg (CC-BY-

SA);http://upload.wikimedia.org/wikipedia/commons/thumb/c/cf/CharaFragilis.jpg/220px-
CharaFragilis.jpg

Habitat

Chara is an epipelic, gregarious alga found attached at the base with the help of root like
rhizoids to the soil at the bottom of stagnant, shallow, slow running, fresh water bodies like
ponds, pools, lakes and streams respectively with aerial parts free floating in submerged

Institute of Lifelong Learning, University of Delhi 2

 Charophyceae

conditions. It is also seen growing in abundance at the banks of Hindon River (Ghaziabad,
UP) in shallow water. The genus has about 117 species of which, 27 are recorded from
India, 7 species are endemic eg. Chara hatei, C. pashanii, C. handae, C. nuda, C wallichii , C
grovesii, and C. vandalurensis.

It grows in hard fresh water with organic matter (producing H2S) Calcium and less O2,
Phosphorus 4-6 mg/l. Chara baltica grows in salt water (salt < 1%). It is an annual or
perennial, hibernates as thick oospores in hot and dry weather and has sulphur containing
compounds due to which it possesses disagreeable onion like odour.

Vegetative Thallus

Figure : Magnified view of Chara Plant (Habit)

Source: Author

Chara thallus is well differentiated into basal root like creamish white rhizoids arising in
bunches(like adventitious roots of higher plant), green or greyish green aerial stem like
axis, surrounded by lateral branches and stipules like leaves. Thallus grows erect in
submerged conditions with jointed appearance of nodes and internodes (Similar to
Equisetum a Pteridophyte) and because of the latter trait is called the Aquatic Horsetail.

It attains a height of 1 meter with profuse lateral growth. C. hatei thallus has a trailing and
subaerial habit and is of 20-30 cm in height. In several species of Chara the thallus bears
rough incrustation of lime as cortical spines that makes it brittle and coarse because of

Institute of Lifelong Learning, University of Delhi 3

 Charophyceae

which it is named as stone wort or brittle wort. Some Species of Chara are flexible and
have mucilage sheath on cell wall, without any other matter deposition.

Rhizoids

A non-green, long, creamish white multicellular anchoring organ. The cells of which have
outer and inner cell wall, nucleus, protoplasm, small solid particles present at the tip of the
rhizoids function as statoliths are involved in graviperception. It arises from the peripheral
cortical cells of lower nodes of main axis, grows with the apical cell and shows oblique
septa.

A B C

Figure: Magnified view of rhizoids : A. An outline diagram showing oblique septum. B-C.
Rhizoidal plate with two basal cells forming rhizoids.

Source: Author

Functions : Rhizoids perform the following functions:

1. Fixing the plant to the soil or substratum.

2. Absorption of minerals from the bottom sediments.
3. Reproduction and perennation by bulbils and secondary protonema.

Institute of Lifelong Learning, University of Delhi 4

 Charophyceae

Figure : Development of rhizoidal initial from a branch of Chara sp. : A-D. Production of 4
lateral cells. E-F. Rhizoids produced only from basal two cells.

Source: Author

Stem

The stem like axis consists of alternate nodes and internodes.

Node: It is a short multicellular and complex structure, which consist of two central cells
surrounded by 6-20 peripheral cells (Cortex).

Figure: Chara - W. M. of main axis with lateral branches

Source: Author

The peripheral cells give rise to two types (dimorphic) of branches:

 Branchelets of limited growth (short)

 Branchelets of unlimited growth (Long)
Internode: It is of two types:

 Corticated internode- An elongated central cell of 10 cm in length surrounded by one

cell thick sheath called cortex eg. C.flaccida. etc.

Institute of Lifelong Learning, University of Delhi 5

 Charophyceae

 Ecorticated internode- Central internodal cell is devoid of peripheral cortical layer of

cells. eg C. nuda. etc

Growth: The main axis or stem grows with the help of a dome shaped apical cell. By means
of transverse walls it cuts disc shaped segments at the posterior or lower end. Each
segment divides further and gives rise to upper biconcave nodal initial and a biconvex
internodal initial. Internodal initial elongates and continue its growth terminally. Nodal initial
divides several times and give rise to 2 central cells and 6 – 20 peripheral cells, secondary
branchlets and bract like stipules or bracts.

Figure: Chara – V.S. of apical metistematic zone.

Source: Author; http://www.biologie.uni-hamburg.de/b-online/fo44/chara50.jpg

The main axis or stem bears two types of lateral branches that arise from nodal region:

Short branches- Branches of limited growth or primary laterals and leaf like structures.
These branches arise in whorls from each node corresponding to the number of peripheral
cells at each node and limit their growth after attaining a definite length. These branches
bear nodes and internodes like the main axis and end up to a long apical cell. At the nodal
region whorl of one celled out growths called bracts or stipuloides or secondary laterals arise
alternating to the branch. The short fertile branches bear sex organs in axil of bracts or
stipuloides, or secondary laterals arise alternate to the branchlets.

Long Branches- They are also named as secondary branches or branches of unlimited
growth. They arise singly or in groups from the axil of primary branchlets; hence are named

Institute of Lifelong Learning, University of Delhi 6

 Charophyceae

as axillary long branches. They have the same structure and growth pattern as of the main
axis. In corticated species these branches cause suppression of upward growth of main axis.

B C

A
D
Figure: Chara – Magnified view of aerial axis showing A. W.M. of branch of limited growth
with fertile lateral branches. B.- C. W.M. appendages at each node. D. Outline diagram of
T.S. internode.

Source: Author

Stipulodes or stipules or bracts: These are short oval or pointed leafy one celled outgrowths
developed from the basal parts of branchlets. The growth pattern shows:

(a) Haplostephanous type with one whorl of bracts on each node, unistipulate (1 bracts)
eg. C. braunii.

(b) Diplostephanous type with two whorl of bracts on each node, Bistipulate (2 bracts)
eg. C. buramanica

(c) Triplostephanous – Stipulodes or bracts occur it three whorls, Tristipulate (3 bracts).

eg. C. ceratophylla

Institute of Lifelong Learning, University of Delhi 7

 Charophyceae

Figure: Chara - Outline diagram of different types of cortex:A.Young cortex with node and
intermodal cells.B.W.M. of haplostichous type with spine cells (stipulodes) C. canescens C-
E.Different stages of diplostichous type C. delicatule F.- H. Triplostichous type C.fragilis

Source: Author

In some species stipules are rudimentary eg. C. nuda or completely absent eg. C. pashanii .

Cortex-Presence of vertically elongated narrow cells around central internodal cell called
cortex. Based on the series of peripheral cortical cells around inter-nodal cells. Cortex is of
three types:-

1. Haplostichous - Single series of cortical cells eg. C. canescence.

2. Diplostichous- Two series of cortical cells eg. C. delicatule.
3. Triplostichous – Three series of cortical cells. eg. C. fragilis.

Cell Structure

Chara has two types of cells:

1. Nodal cell:- It is a small isodiametric in outline with single centrally placed nucleus and
small scattered vacuoles.

Institute of Lifelong Learning, University of Delhi 8

 Charophyceae

2. Internodal cell – It is long and cylindrical. The adjacent cell exhibits protoplasmic
continuity by means of plasmodestmata. Each cell is surrounded by:

 Outer cell wall which is made up of cellulose (found in the form of microfibrils
deposited in a homogeneous matrix) .

 Inner cell membrane hyaline, selectively permeable.

 A superficial layer of mucilaginous material in some species.

 Lime in the form of calcite (Rhomboidal) deposits on the cell wall in several
species.

Protoplasm contains cytoplasm, endoplasmic reticulum, mitochondria, ribosomes,

dictyosomes, chloroplasts, nuclei and large central vacuole. The outer exoplasm has
partially embedded chloroplasts. The inner endoplasm is more fluid than the outer
exoplasm. Pyrenoids are absent. Reserve food material is in the form of starch grains. The
internodal cell is multinucleated The number of chromosomes vary eg. n=12 in C.
canescense and n=28 in C. zeylanica.

Figure: Chara- Magnified view of a chloroplast

Source: Author

Institute of Lifelong Learning, University of Delhi 9

 Charophyceae

A. B.

Figure: Chara sp. A. Magnified view of internodal cell. B. Protoplasmic streaming in a

internodal cell.

Source: Author

Streaming of Protoplasm (Cyclosis): All types of cells except young show cyclosis. The
endoplasm of plasmasol continually show up and down streaming in long direction, which is
caused by sol-gel changes in the cytoplasm through alternate contraction and expansion of
protein fibrils.

Reproduction

Asexual Reproduction

Chara reproduces asexually by vegetative means. The vegetative parts of the plant give rise
to small protuberances in hot and dry weather. Asexual spores are absent.

Institute of Lifelong Learning, University of Delhi 10

 Charophyceae

Figure: Chara W. M. of asexual (veg.) reproductive bodies. A. Amylum stars, B. Root

bulbs, C. Amorphous bulbils

Source: Authors

 Tubers and Bulbs : The small outgrowths that arise from the lower nodes and rhizoids.
These outgrowths are oval or spherical in shape and rich in food eg C. aspera.

 Amorphous and spherical bulbils: These are aggregated small and swollen cells that
are formed by irregular division of all peripheral cells of stem or all the four cells of the
rhizoidal joint. Tubers and Bulbils perennate during unfavourable conditions and grow in
next favourable season to form new plants.

 Amylum Stars: The multicellular star shaped aggregates develop by proliferation of

nodal cells at lower nodes and are filled with abundant amylum starch as reserve food.
These stars when detached can give rise to a new plant. eg. C. stelligera.

 Secondary Protonema: The protonemal structures arise from several parts of the plant
which later give rise to adult plants:

 Rhizoidal node of Primary protonema.

 Basal node of Primary rhizoids
 Dormant apical cells.
 Nodal cells of axis or stem.

 Buried nodes of older plants

Sexual Reproduction
Sexual reproduction is oogamous. In submerged habitats when temperature rises, water
recedes, minerals increase and pH changes (drying up conditions), these conditions hasten

Institute of Lifelong Learning, University of Delhi 11

 Charophyceae

the formation of sex organs. The well-differentiated complex and macroscopic antheridium
and oogonium are borne on the adaxial surface of lower branchlet nodes.

Figure: Chara sp. Magnified view of a fertile branch to show the position and details of
nucule and globule.

Source: http://upload.wikimedia.org/wikipedia/commons/2/26/CharaV3.jpg;Author;
http://facultyweb.berry.edu/mcipollini/bio311/files/chara.jpg

Majority of species are monoecious or homothallic and bear both the sex organs on the
same thallus eg. C. zelanica. A few species are dioecious or heterothallic and bear both the
sex organs on different thalli eg. C. wallichi etc. In monoecious species the oogonium lies
above the antheridium at the same branchlet node. In some species the monoecious habit
changes to dioecious habit by change in day length eg. C corallina. Sex organs are
considered to be condensed branches and are quite similar to sex organs of bryophytes .

Antheridium or globule

Institute of Lifelong Learning, University of Delhi 12

 Charophyceae

It is a globose, green at young stage and turns yellow, orange or reddish in colour towards
maturity. It is borne at the top of a branch of second order and replaces a bract. It has a
large pedicel cell lying partly embedded inside the body of antheridium. The basal node
contains five peripheral cells. Two of the lateral peripheral cells produce one celled
bracteoles, the lower two peripheral cells grow into cortical filaments in corticated forms. In
monoecious species the upper peripheral cell gives rise to oogonium or nucule.

Figure: Chara- A-F. Developmental stages of globule from apical lateral cell; G- Whole
mount of mature globule; H. V.L.S. of mature globule; I. W.M. of sheild cell, Manubrium
and Anthredial filaments; J. Magnified view of spermatogenous cells; K. Single antherozoid;
L. W.M. of Antherozoid (sperm).

Source: Author

Structure

 Shield Cell: These are concave or convex plate like eight large curved cells initially green
due to chloroplast pigments which later turns brilliant orange coloured due to gamma
carotene in abundance. These cells have large number of wall infoldings which show
sutures fitting into one another.

 Manubrium: It is eight in number and project towards the centre of the globule.
Manubrium arises from the central part of the shield cell as an elongated club-shaped

Institute of Lifelong Learning, University of Delhi 13

 Charophyceae

cell. Each manubrium bears a head cell or primary capitulum. The cell cuts secondary
capitular cells which further divide 2-4 times to form terminal cell initials. These
terminal cell initials divide mitotically to give rise to antherozoidal branched or
unbranched filament at the centre called spermatogenous filaments.

 Spermatogenous filament or an antheridial filament consists of series of cells arranged

in uniseriate fashion called antherozoid mother cells or sperm mother cells. Each cell of
spermatogenous filament metamorphoses to give rise to biflagellate sperms or motile
male gametes (n). Each globule or antheridium produces two thousand or more sperms.

 Spermatozoid: The sperm or a male gamete is narrow, elongate, spirally coiled

uninucleate, unicellular, hyaline, biflagellated structure that has the cytoplasm enclosed
at the posterior end. The body shows three coils or turns, the two flagella arise
subterminally at the anterior end and are smooth and similar type.

 With the maturity of sperms, the shield cells separate due to internal pressure of
manubria, capitular cells and enzymatic secretions. The sperms come out of the pore
with body first and flagella at the end in the early hours of morning and continue to
swim to reach the nucule slit till evening.

Oogonium or Nucule

It is a large ellipsoidal, oblong single celled female sex organ which encloses a single
gamete or egg and is borne on the top of four celled filament. The three cells lying below
the nucule are the stalk cell, nodal cell and the pedicel cell. The pedicel is connected
adaxially (dorsally) with basal nodal cell of the antheridial axis (in monoecious forms) and
branchlet node (in dioceous forms). The nodal cell produces five peripheral envelope cells
which elongate and spread over the oogonium. These cells divide transversely into upper
coronular cells (corona) and lower tube cells. The tube cells get twisted spirally in a
clockwise (right to left from below) direction and are called spiral cell bracts. The
ensheathing cells are green when young but after fertilization they become yellow or brown.

The twisting of spiral cells serves two functions:

 Keeping contact with oogonium.

 Exudation of chemotactic mucilaginous ooze at the time of fertilization.

Institute of Lifelong Learning, University of Delhi 14

 Charophyceae

Oogonium is a one celled female sex organ enclosing a single female non-motile haploid egg
or ovum(n). It has a thin wall, single nucleus lying at the base, chloroplast, abundant starch
grains, little cytoplasm and a receptive spot for the entry of the serms.

Fertilization : At maturity, the receptive spot or area in the apical part exudes
mucilaginous (ooze) fluid. Tube cells swell below corona, elongate and separate terminally
from one another to create five slit opening. Mucilage oozes out and attracts the sperms by
chemotactic influence. Several sperms enter the receptive spot but only one penetrates the
oosphere or ovum or female gamete and fuse to produce diploid zygote.

 Zygote or Oospore : During maturation of zygote the envelope cells turn yellowish
brown and store a lot of lime minerals. Inner walls of tube cells thicken, suberize and
silicify. Oogonial wall also thickens, develops exine (outer thick layer) , mesine (middle
layer) and intine (inner layer) to enclose diploid nucleus (2n) and cytoplasm. Reserve
food starch gets converted into fat (oil drops) and the oospore becomes dark brown,
seed like structure. The resting oospore settles down on the soil or sand of stagnant
water to perennate after the degradation of plant body. It is a thick walled unicellular,
uninucleated diploid (2n), hard , ovate , ellipsoidal in outline (eg. C nuda), dark brown or
golden brown with spiral ridges on its surface. It is a perennating structure resembling
the seeds of higher plants.

Figure: Chara- A. Developmental stages of nucule from apical oogonial cell.; B. V.S. of
nucule showing internal structures. C. W.M. of nucule with primary, secondary and
tertiary bracts like angiospermous female flower.

Institute of Lifelong Learning, University of Delhi 15

 Charophyceae

Sourec: Author

A. B.

Figure: A- B. Chara – A photograph of nucule ( orangish in color) resembles the

angiospermous flower.

Source:http://postfiles7.naver.net/20100125_54/nstdaily_12644283647422iYCi_jpg/%EC%B0%A8%EC%B6%9
5%EC%A1%B0%28chara_braunii%29_-
_%EC%9C%A4%EC%A1%B0%EB%A5%98,_%EC%9C%A4%EC%A1%B0%EC%8B%9D%EB%AC%BC_nstdaily.jp
g?type=w3;
http://postfiles10.naver.net/20100125_217/nstdaily_1264428368661UjRH9_jpg/%EC%B0%A8%EC%B6%95%EC
%A1%B0%28chara_braunii%29_-
_%EC%9C%A4%EC%A1%B0%EB%A5%98,_%EC%9C%A4%EC%A1%B0%EC%8B%9D%EB%AC%BC4_nstdaily.j
pg?type=w3

Figure: Chara- A. Spermatozoids near ostiole at the coronary ostiole. B. V.S. of

developing oospore (2n) enclosed by oospore wall. C. W.M. of mature oospore or seed (2n)
showing ridged exine. D. W.M. of ridges seen at the base of the oospore.

Source: Author

Institute of Lifelong Learning, University of Delhi 16

 Charophyceae

Figure: C. fragilis – A photograph of W. M. of zygote (2n) enclosed in nucule.

Source: http://mcgregor.sbs.auckland.ac.nz/files/2011/01/dave_082_1.jpg

Germination of Oospore or zygote : The three membranous thick walled diploid oospore
germinates during favourable conditions on sand or soil in submerged habitat with the
following conditions:

 sufficient light (towards red side)

 temperature 20-27±1oC
 pH 5-10
 minerals like Phosphorous 20-40 mg/l
 Less oxygen, calcium
 rich organic matter.
Oospore is a thick walled seed like structure that absorbs water, minerals and germinates.
The outermost exine - thick dark brown inelastic layer cracks open by imbibition and the
mesine, intine (inner layers) along with protoplasm and nucleus exude out as a small
germling or germ tube at the apical tip. The diploid nucleus (2n) undergoes meiosis to form
four haploid nuclei (n). A curved wall appears at the apical part. It divides the interior of the
oospore into two unequal halves (cells), the smaller lenticular apical cell and the larger
basal cell. Apical cell divides longitudinally into two cells, the protonemal initial and the
rhizoidal initial. Both these cells grow out to form erect filamentous aerial primary
protonema and basal primary rhizoids. This later gives rise to the Chara plant. These stages

Institute of Lifelong Learning, University of Delhi 17

 Charophyceae

resemble with Batrachospermum a red alga. Protonemal stage is also found in some
bryophytes.

Figure: Chara- Different stages of germination of oospore shown in vertical section of

oospore. A. A resting zygote; B. Meiosis; C. Cracking of outer exine and emergence of
germling. D. Separation of upper protonemal and lower rhizoidal initial. E. Growth of
primary protonemal and rhizoidal initials.; F. Young protonemal aerial plant with lower
primary rhizoids.

Source: Author

Institute of Lifelong Learning, University of Delhi 18

 Charophyceae

Figure: Chara- Young protonemal plants with aerial branching and secondary protonemal
branches arising from lower rhizoidal node.

Source: Author

Figure: Chara – An outline view of life cycle.

Source: Author

Also visit: http://physwiki.apps01.yorku.ca/images/thumb/5/51/02_Figure_2.5.PNG/400px-

02_Figure_2.5.PNG

Institute of Lifelong Learning, University of Delhi 19

 Charophyceae

Parthenogenesis- This process is seen in nature in some species of Chara.The nucule or

oogonium directly produces thick wall resting oospores without fertilization.These spores or
parthenospores directly germinate in favourable conditions and produce new plants. This
process is common in dioecious species like C. canescens.

Economic importance
Charales exhibit the dominant forms of macrophytic green algae in base rich fresh waters
for some three million years earlier than embryophytes. They are important in many
respects with regard to food, calcification, ecology and as experimental tools:

 Charales are used as food for aquatic animals.

 Chara plant supports many epiphytic herbivores being used as food for fishes.

 Bulbils are eaten by diving birds or water fowls.

 Growth of Chara can clear turbid water. Dried plants possess pungent odour i.e.
being used as insect repellent in farmhouses.

 Charales have larvaecidal property.

 Lime deposits at the bottom of water reservoirs appear as marl and tufa. These are
used in the manufacture of cement and also for soil reconditioning.

 Charales are used as manure and also to correct the acidity of soil.

 These are used as fertilizer as they form good crumb structure and aeration to
clayey soil.

 Charales because of macroscopic size are a favorable material for physiologists to

carry out experimental studies of:

Electrophysiological features of membrane functions.

 Analysis of cytoplasmic streaming and geotropism.

 Charales due to their abundant growth in water reservoirs disrupt the proper working
or supply of water.

Institute of Lifelong Learning, University of Delhi 20

 Charophyceae

Systematic Position and Relationship with other green

plants

Charales have been considered an advanced complex, modified group amongst

Chlorophyceae based on micromorphological, ultrastructure, biochemical, molecular and
physiological features. Phycologists proposed a distinct position of Charales at the rank of
division, class under Thallophytes.

Fritsch 1935 and Iyenger 1951 considered Charales at an order level of Chlorophyceae
(green algae) based on morphological and reproductive features

 Bold 1967 classified green algae to Phyllum- Chlorophycophycophyta and the order
Charales was separated and raised to the rank of Phyllum – Charophyta

 Desikachari and Sudaralingum 1962, Round 1973, and other phycologists preferred to
place the stoneworts in Division – Charophyta separate and parallel to Chlorophyta.

The Characters which place Charalean algae in or near green algae:-

 Presence of green pigment chlorophyll a and b.

 Reserve food material – starch
 Haploid nature of plant body.
The features which characterize Charales as distinct group:

 Erect vegetative plant body with distinct nodes and internodes.

 Dimorphic whorled lateral branches.

 Cortication and occurrence of sex organs on the lateral branches.

These characters have formed the basis of the origin of Charales from Chaetophorales (eg.
Draparnaldiopsis) or their ancestors.

Chaetophorales Charales.

Charalean group exhibits distinct characters which brings it closer to the other plant groups
– Bryophytes and Pteridophytes:

 Vegetative growth by apical cell.

Institute of Lifelong Learning, University of Delhi 21

 Charophyceae

 Young protonemal stage with the germination of zygote or oospore (2n) after
meiosis.
 Rhizoids uniseriate, multicellular with oblique septa.
 Stem with knotted nodes and internodes (similar to Equisetum) and peculiar mode of
branching.
 Sex organs multicellular and their developmental stages.
 A layer of sterile tissue present around the sex organs.
 The tube cells or corona of nucule or oogonium resembles the venter and neck cells
of archegonium in bryophytes.
 Spirally coiled antherozoids with flagella having band of micro-tubules at their base.

Charalean green algae show affinities with embryophytes :

 Presence of discoid chloroplast.
 Multicellular complex plant body with stem like axis, root like rhizoids, leaf like
stipulodes or bracts.
 Meristematic growth by apical cell.
 Asexual reproduction by vegetative structures like tubers, bulbils and protonema.
 Sexual reproduction by multicellular male gametophytic globule enclosing male
unicellular motile gametes (n), and female gametangial oogonium enclosing egg or
ovum (n) surrounded by three membranous structures like ovary and externally by
primary, secondary bracts similar to the calyx lobes and corona like corolla of
flowering plants.
 Zygote or oospore resembles a thick walled nut or seed like hibernating structure.
 Phragmoplasts (persistent spindle and glycolate oxidase) have been seen in Charales
, Coleochaetales and Zygnemales (Spirogyra). In the dividing cells the telophasic
nuclei lie far apart due to open spindle formation.
 Primary plasmodesmata (during cytokinesis) develop much like those of land plants
and also is seen in one of the Chara species.
 The developmental and morphological features thus reveal that Charales and
Coleochaetales are closely associated with embryophytes.

The Charales and Coleochaetales are considered as the closest living green
algal relatives of land plants

Institute of Lifelong Learning, University of Delhi 22

 Charophyceae

Source:http://tolweb.org/Public/treeImages/Green_plants.png?x=1203662046

The green plants ( assemblage of photosynthetic organisms, containing chlorophyll a

and b, having double membrane bound chloroplasts and cellulosic cell walls) are
divided into two main lineages:

 Green algal lineage that includes the Prasinophytes, Chlorophyceae,

Trebouxiophyceae and Ulvophyceae.
 Streptophytes
Traditionally Charales and Coleochaetales were placed under Charophyceae, Division
Chlorophyta. However ultrastructural, morphological as well as molecular studies
support the fact that Charophyceae is a paraphyletic group and are most closely
related to higher land plants. These orders therefore have been now placed within
the Streptophyta

Lee 2008 grouped algae into four distinct groups (gene sequence, ultra structure of
organalles and other molecular features taken into account )
 Prokaryotes –Cyanophyta (Cyanobacteria ).
 Eukaryotic Algae (chloroplast surrounded by two membranes)
 Eukaryotic Algae with chloroplast surrounded by one membrane of chloroplast
endoplasmic reticulum.
 Eukaryotic Algae with chloroplast surrounded by two membrane of chloroplast
endoplasmic reticulum.
Under this system Charophyceae occupies distinct position as a class under Group 2
Division Chlorophyta.

Institute of Lifelong Learning, University of Delhi 23

 Charophyceae

Systematic Position
Fritsch (1935) Bold and Wynne (1978) Lee( 2008)
Division – Algae Division - Charophyta Phylum - Chlorophyta

Class – Chlorophyceae Class - Charophyceae Class - Charophyceae

Order – Charales Order - Charales Order - Charales

Family- Characeae Family - Characeae Family - Characeae

Genus –Chara Genus - Chara Genus - Chara

Summary

Chara sp. is a highly complex, well defined green alga. Its thallus is differentiated into root
like rhizoids, stem like aerial axis and leaf like lateral unicellular stipulodes or bracts. It
reproduces asexually by vegetative means and sexually by oogamy. This alga shows
affinities with bryophytes, pteridophytes and embryophytes. Chara plant body and
reproductive structures justify its placement as a distinct group - Charophyta at division
level parallel to Chlorophyta. Charales (Chara and Nitella ) the largest and complex group
amongst Chlorophyceae with their plant body reaching one metre length along with whorls
of lateral branches at each node, show the great similarity with aquatic flowering plants (eg.
Ceratophyllum sp.) in submerged conditions. In addition, the recent studies with regard to
the features like molecular sequence, architectural evidences that include intron insertion
steps and movement of genes between genomes suggest that the order zygnemales of
Chlorophyceae originated prior to Chaetophorales, Charales (green algae) and
embryophytes (multicellular flowering plants). These features strongly represent the
monophyletic relationship of this algal group with embryophytes and supports the fact that
their ancestors do share features with members of Charalean green algae.

Exercises

1. Comment on the following:

a) Germination of oospore in Chara
b) Epipelic nature of stone worts.
c) Charales resemble Chlorophyceae.
2. Write short notes on the following:
a) Nucule.

Institute of Lifelong Learning, University of Delhi 24

 Charophyceae

b) Globule
c) A sexual reproduction in Chara.
3. Fill in the blanks.
a) Pyrenoids are------------ in chara.
b) Reserve food material of chara ------------.
c) Stone worts is a name of------.
d) Protoplasmic streaming in a cell is called -------.
e) Male sex organs in Chara is called-----.
f) Oogonium of chara is called---------.
g) Root like anchoring organ in Chara------.
4. Draw well labelled diagrams:
a. W.M of sex organs of Chara.
b. V.L.S of Globule.

Glossary

Antheridium : A cell that undergoes internal division and for differentiation to form male
gamete.

Apical cell(s) : The single apical cell or group of cells that occur at the tip of a thallus,
often capable of merismatic proliferation (apical growth).

Benthic: Growing in the benthes – the bottom of a lake, stream or marine system.

Calcification: Deposition of calcium carbonate.

Cell wall: A more or less rigid enclosure surrounding the cell membrane; can be composed
of various materials.

Cellulose: A β14 linked glucose that forms micro fibrils in the cell walls of various algae
and all land plants.

Embryophytes: Bryophytes and vascular plants.

Epipelics: Living on the surfaces of mud and sand (Rooted at the base and free floating
arial parts in submerged conditions).

Gyrogonites: The calcified zygotes of Charalean green algae; includes those occurring in
the fossil record and those in modern sediments.

Institute of Lifelong Learning, University of Delhi 25

 Charophyceae

Internodal cell: A cell located between nodal cells.

Brackish : Saline water with a salinity less than that of see water.

Manubria (sing.Manubrium):In Charalean green algae; columnar cells that connect the
pedicel to the shield cells within male gametangia.

Oogamy: Sexual reproduction involving syngamy of a small flagellate male gamete and a
larger, nonflagellate female gamete.

Oogonium: A structure that produces one or more eggs.

Parthenogenesis: Production of a new individual from an unfertilized gamete.

Plamasol: Protoplasm

Shield cells: In Charalean green algae, cells that form the outer walls of the multicellular
male gametangia.

Spermatogenous threads: In Charalean green algae, filaments of cells that each produce
a sperm, located within a complex antheridium.

Sporopollenin: In walls of certain green algae and spores of embryophytes a resistant

biopolymer.

Systematics: The scientific study of organismal diversity and the relationships among
organisms.

Taxon: A general term for any taxonomic category.

Tube cells: In Charalean green algae, elongated helically twisted tubular cells that
surround oogonia.

Zygote: The product of gamete fusion (syngamy).

Amylum star: Star shaped aggregate of cells filled with amylum starch that forms new
plants in the stoneworts (charales)

Plasma membrane: (Plasmalemma) The outermost living membrane of a cell.

Plasmodesmata : The minute cytoplasmic threads that extend through openings in cell
walls and connect the protoplasts of adjacent living cells.

Institute of Lifelong Learning, University of Delhi 26

 Charophyceae

Spermatogenesis: Formation of sperm.

Sperm: Male gamete.

References/ Suggested Readings

Bold, H.C. 1967. Morphology of Plants. Harper and Row, New York.

Desikachary, T.V. and Sundaralingam, V.S. 1962. Affinities and interrelationships of the
Characeae. Phycologia, 2:9-16.

Fritsch, F.E.1935. The Structure and Reproduction of Algae, Vol. 1 & 2. Cambridge
University Press.

Bold, H.C. and Wynne, M.J. 1985. Introduction to the Algae: Structure and Reproduction,
Second Ed. Prentice Hall, New Jersey.

Bell, P.R and Woodcock, L.F Diversity of Green plants.

Bhatia, K.N 1993, A treatise on Algae, R. Chand and Co.

Round, F.E. 1973. The Biology of Algae, 2nd Edition. St. Martin Press, New, York.

Borowitzke M.A, 1982 Morphological and cytological aspects of algal calcification Int. Rev.
Cytol. 74 : 127-62.

Casanova, MT. 1997 Oospore variation in three species of chara (charales, chlorophyte).
Phycologia, 36 : 274-80.

Goutchasov, A.A, Marin, B & Melkonian, M 2002 Molecular Phylogeny of conjugating

green Algae (zygnemophyceae, streptophyta ) inferred from SSU DNA sequence comparison
J.Mol. Evol. 56 : 89-104.

Graham L.E. 1993, The origin of land plants. John Wiley, New York, NY.

Graham, I.E 1984 coleochaete and the origin of Land plants Am.J.Bot 71 : 603- 608.

Graham, L.E and Wilcox, L.W 2000, Algae, Prentice Hall Inc.

Hodick . D. 1994 Negative gravitropism in chara protonemata a model integrating the

opposite graitropic responses of protonemate and rhizoides. Planta, 195 : 43-49.

Kenrick, P and Crane, P.R 1997, The Origin and Early Diversification of land plants.
Smithsonion Institution Press, Washinton, DC.

Institute of Lifelong Learning, University of Delhi 27

 Charophyceae

Kwiatkowske, M 2003 Plasmodesmal changes are related to different developmental stages

of antheridia of chara species. Protoplasma, 222 : 1-11.

Lee, R.E. 2008, Phycology, Cambridge univ. Press 4th Edition.

Levis I.A & Mc Court, R.M. 2004, Green Algae & the origin of land plants . Amer. J.Bot 91 :
1535-56.

Lucas W.C. 1979 Alkaline band formation in chara coralline. Plt.Physiol 63 : 248-54

Piickett- Heaps, J.D 1975. Green Algae. Sinauer Associates, Sunderlands, MA

Links

http://www.biologie.uni-hamburg.de/b-online/e44/chara.htm

http://taibif.org.tw/nbrpp/algae.php?select=Lychnothamnus+barbatus

http://aquaplant.tamu.edu/plant-identification/visual-index/muskgrass/

http://comenius.susqu.edu/biol/202/archaeplastida/viridiplantae/green%20algae/charophyt
a/charophyta.htm

Institute of Lifelong Learning, University of Delhi 28