How Did Plants Evolve?

Red algae and green algae are the

closest relatives of land plants

 Over a billion years ago, a heterotrophic protist acquired a

cyanobacterial endosymbiont
 The photosynthetic descendants of this ancient protist evolved into red
algae and green algae

1 2

Red Algae
Algae Are Protists
 Red algae are reddish in color due to an accessory pigment call
Diplomonads Diplomonadida

Chlorophyta
Animalia

phycoerythrin, which masks the green of chlorophyll

Rhodophyta
Euglenozoa
Parabasalids Parabasala

Plantae
Radiolarians Radiolaria
Cercozoa

Fungi

 Red algae are usually multicellular; the largest are seaweeds

Alveolata Stramenopila Amoebozoa (Opisthokonta) (Viridiplantae)
 Red algae are the most abundant large algae in coastal waters of the
Choanoflagellates
Dinoflagellates

Entamoebas
Chlorarachniophytes
Ciliates

Plants
Kinetoplastids

Euglenids

Charophyceans
Oomycetes

Metazoans
Apicomplexans

Foraminiferans

Gymnamoebas
Diatoms

Chlorophytes
Cellular slime molds
Plasmodial slime molds

Fungi
Golden algae

Red algae
Brown algae

tropics

Ancestral eukaryote

3 4

LE 28-28
Green Algae

Dulse (Palmaria palmata). This  Green algae are named for their grass-green chloroplasts
edible species has a “leafy” form.
 Two main groups: chlorophytes and charophyceans (closest relatives of
land plants—will get into next)
 They are closely related to land plants

A coralline alga. The cells walls of

corralline algae are hardened by
calcium carbonate. Some coralline
Bonnemaisonia hamifera, a algae are members of the biological
filamentous red alga. communities called coral reefs.

5 6

1
 Overview: The Greening of Earth

 M ost chlorophytes live in fresh water, although many are marine

 Looking at a lush landscape, it is difficult to imagine the land without
 Other chlorophytes live in damp soil, as symbionts in lichens, or in snow any plants or other organisms
 Chlorophytes include unicellular, colonial, and multicellular forms  For more than the first 3 billion years of Earth’s history, the terrestrial
surface was lifeless
 Since colonizing land, plants have diversified into roughly 290,000 living
species

7 8

LE 29-3
Land plants evolved from green algae

 Green algae called charophyceans are the closest relatives of land

plants
 Comparisons of both nuclear and chloroplast genes point to
charophyceans as the closest living relatives of land plants

Chara, 10 mm
a pond
organism
(LM).

40 µm

Coleochaete orbicularis, a disk-shaped

charophycean (LM).

9 10

The Origin and Diversification of Plants

Diplomonads Diplomonadida

Chlorophyta

 Fossil evidence indicates that plants were on land at least 475 million
Animalia

Rhodophyta
Euglenozoa
Parabasalids Parabasala

Plantae
Radiolarians Radiolaria

years ago
Cercozoa

Fungi

Alveolata Stramenopila Amoebozoa (Opisthokonta) (Viridiplantae)  Fossilized spores and tissues have been extracted from 475-million-year-
old rocks
Dinoflagellates
Kinetoplastids

Choanoflagellates
Entamoebas
Oomycetes

Chlorarachniophytes
Ciliates

Plants
Euglenids

Charophyceans
Metazoans
Apicomplexans

Foraminiferans

Gymnamoebas
Diatoms

Chlorophytes
Cellular slime molds
Plasmodial slime molds

Fungi

Red algae
Golden algae
Brown algae

Ancestral eukaryote

11 12

2
 How to Deal? Evolutionary
Adapting to life on land Innovations…
 What are some of the challenges that plants would have to contend
with if they moved onto land?  Cuticle, a waxy covering over leaves, cut down on water loss through
transpiration and allowed early land plants to tolerate drying
 Vascular tissues permitted land plants to grow beyond small size. These
cells provide a mechanism for conducting water along the potential
-Light (too much or too little?) gradient created by uptake at the roots and loss at the leaves. This
process is much more efficient than osmosis.
-Water  Pollen and seeds are multicelluar structures with a protective coat, and
resist drying out much better than sperm and spores.
-Nutrients

Therefore, many adaptations emerged after land

plants diverged from their charophycean
relatives

13 14

Vascular vs. Non-Vascular

 Those ancestral species gave rise to a vast diversity of modern plants Seed Plants Seedless Bryophytes
Land plants can be informally grouped
Lycophyta (club/spike moss, Liverworts
based on the presence or absence of quillworts) Hornworts
Pterophyta (ferns, horsetails, Mosses
vascular tissue whisk ferns)

Gymnosperms Angiosperms
(naked seeds) (container/fruit seeds)

Coniferophyta (Conifers)
Monocots Dicots
Cyadophyta (Cyads—palm-like)
Ginkophyta (Ginkos)
Gnetophyta

15 16

LE 29-7
Land plants The life cycles of mosses and other bryophytes are
Vascular plants
dominated by the gametophyte stage
Bryophytes Seedless vascular plants Seed plants

 Bryophytes are represented today by three phyla of small herbaceous

(nonwoody) plants:
sperms
Gymno-
Pterophytes
Hornworts

Mosses

Lycophytes
Liverworts

sperms
Angio-
Charophyceans

 Liverworts, phylum Hepatophyta

 Hornworts, phylum Anthocerophyta
Origin of seed plants
(about 360 mya)  Mosses, phylum Bryophyta

Origin of vascular plants

 M osses are most closely related to vascular plants
(about 420 mya)

Origin of land plants

(about 475 mya)

Ancestral
green alga

17 18

3
 Alteration of Generations Gametophyte
Makes the gametes/sex cells that
combine to make a moss plant

 Like all plants, mosses undergo alteration of generations

 Alteration of Generations is a life cycle in which there is both a
multicellular diploid form (the sporophyte), and a multicellular haploid
form (the gametophyte)—characteristic of plants and some algae.
 In mosses and other bryophytes, it’s the gametophyte that is the
dominant form of the plant.

19 20

LE 29-5b LE 29-9b
Alternation of Generations

Haploid multicellular
organism (gametophyte) Plagiochila
Mitosis Mitosis deltoidea,
a “leafy”
liverwort
Spores Gametes

MEIOSIS FERTILIZATION

Zygote

Mitosis
Diploid multicellular
organism (sporophyte)

21 22

Vascular vs. Non-Vascular

Moss
Polytrichum
commune,
hairy cap
moss Seed Plants Seedless Bryophytes
Sporophyte
Lycophyta (club/spike moss, Liverworts
quillworts) Hornworts
Pterophyta (ferns, horsetails, Mosses
whisk ferns)

Gymnosperms Angiosperms
(naked seeds) (container/fruit seeds)

Gametophyte Coniferophyta (Conifers)

Monocots Dicots
Cyadophyta (Cyads—palm-like)
Ginkophyta (Ginkos)
Gnetophyta

23 24

4
 Ferns and other seedless vascular plants Origins and Traits of Vascular Plants
formed the first forests
 Bryophytes and bryophyte-like plants were the prevalent vegetation  Fossils of the forerunners of vascular plants date back about 420 million
during the first 100 million years of plant evolution years

 Vascular plants began to diversify during the Carboniferous period  These early tiny plants had independent, branching sporophytes

 Vascular plants dominate most landscapes today  They lacked other derived traits of vascular plants

25 26

Life Cycles with Dominant Sporophytes

Key
Haploid (n)
Diploid (2n)
 In contrast with bryophytes, sporophytes of seedless vascular plants are
the larger generation, as in the familiar leafy fern Spore Young Antheridium
gametophyte
 So, in the earliest land plants (bryophytes), the spore-producing MEIOSIS

organism was larger than the gamete producing organism. In vascular

seedless plants (like ferns), it’s the spore-producing organism that’s the Sporangium
Sperm
largest. (The gametophyte is much smaller and harder to see.)
Archegonium
 The gametophytes are tiny plants that grow on or below the soil surface Egg

Sporangium Mature New

sporophyte sporophyte Zygote FERTILIZATION
Sorus

Gametophyte

Fiddlehead

No male/female gametophytes in fern spores—they are bisexual/homospores

27 28

Transport in Xylem and Phloem Evolution of Roots

 Vascular plants have two types of vascular tissue: xylem and phloem  Roots are organs that anchor vascular plants
 Xylem conducts most of the water and minerals and includes dead  They enable vascular plants to absorb water and nutrients from the soil
cells called tracheids
 Roots may have evolved from subterranean stems
 Phloem consists of living cells and distributes sugars, amino acids, and
other organic products

29 30

5
 Evolution of Leaves

 Leaves are organs that increase the surface area of vascular plants,
thereby capturing more solar energy that is used for photosynthesis
 Leaves are categorized by two types:
 Microphylls, leaves with a single vein
 Megaphylls, leaves with a highly branched vascular system
 According to one model of evolution, microphylls evolved first, as
outgrowths of stems

31 32

LE 29-13
Vascular tissue
Classification of Seedless Vascular Plants

 There are two phyla of seedless vascular plants:

 Lycophyta includes club mosses, spike mosses, and quillworts
 Pterophyta includes ferns, horsetails, and whisk ferns and their relatives
Microphylls

Megaphylls

33 34

LE 29-14a LE 29-14b
Selaginella apoda,
a spike moss

Isoetes
gunnii,
a quillwort

35 36

6
 Phylum Pterophyta: Ferns, Horsetails, and Whisk
Ferns and Relatives
• Ferns are the most diverse seedless vascular
plants, with more than 12,000 species
• They are most diverse in the tropics but also
thrive in temperate forests
The Significance of Seedless Vascular
Plants
 The ancestors of modern lycophytes, horsetails, and ferns grew to great
heights during the Carboniferous, forming the first forests
 These forests may have helped produce the global cooling at the end
of the Carboniferous period
 The decaying plants of these Carboniferous forests eventually became
coal

• Some species are even adapted to arid

climates

37 38

Vascular vs. Non-Vascular

Seed Plants Seedless Bryophytes

Lycophyta (club/spike moss, Liverworts

quillworts) Hornworts
Pterophyta (ferns, horsetails, Mosses
whisk ferns)

Gymnosperms Angiosperms
(naked seeds) (container/fruit seeds)

Coniferophyta (Conifers)
Monocots Dicots
Cyadophyta (Cyads—palm-like)
Ginkophyta (Ginkos)
Gnetophyta

39 40

Plant Diversity II: The Overview: Feeding the World

Evolution of Seed Plants  Seeds changed the course of plant evolution, enabling their bearers to
become the dominant producers in most terrestrial ecosystems

41 42

7
 The Evolutionary Advantage of Seeds Gymnosperms bear “naked” seeds,
typically on cones
 A seed develops from the whole ovule  The gymnosperms include four phyla:
 A seed is a sporophyte embryo, along with its food supply, packaged in  Cycadophyta (cycads)
a protective coat  Gingkophyta (one living species: Ginkgo biloba)
 Gymnosperms (like pines) have “naked seeds,” where the seed is not  Gnetophyta (three genera: Gnetum, Ephedra, Welwitschia)
enclosed in a protective ovary—think pine nut
 Coniferophyta (conifers, such as pine, fir, and redwood)
 Angiosperms have seeds that are protected by an ovary—think apple or
sunflower seed

43 44

LE 30-4aa LE 30-4ac

Cycas revoluta

45 46

LE 30-4ae LE 30-4bc
Bristlecone pine.
Ephedra. This genus includes about 40 species that inhabit arid This species (Pinus
regions throughout the world. Known in North America as longaeva), which is
“Mormon tea,” these desert shrubs produce the compound found in the White
ephedrine, commonly used as a decongestant. Mountains of
California, includes
some of the oldest
living organisms,
reaching ages of
more than 4,600
years. One tree (not
shown here) is
called Methuselah
because it may be
the world’s oldest
living tree. In order
to protect the tree,
scientists keep its
location a secret.

47 48

8
 Vascular vs. Non-Vascular
LE 30-4bd
Sequoia. This giant
sequoia (Sequoiadendron
giganteum), in California’s
Sequoia National Park
weighs about 2,500 metric Seed Plants Seedless Bryophytes
tons, equivalent to about
24 blue whales (the
largest animals), or 40,000 Lycophyta (club/spike moss, Liverworts
people. Giant sequoias quillworts) Hornworts
are the largest living
organisms and also some Pterophyta (ferns, horsetails, Mosses
of the most ancient, with whisk ferns)
some estimated to be
between 1,800 and 2,700
years old. Their cousins, Gymnosperms Angiosperms
the coast redwoods (naked seeds) (container/fruit seeds)
(Sequoia sempervirens),
grow to heights of more
than 110 meters (taller
than the Statue of Liberty) Coniferophyta (Conifers)
Monocots Dicots
and are found only in a Cyadophyta (Cyads—palm-like)
narrow coastal strip of Ginkophyta (Ginkos)
northern California. Gnetophyta

49 50

Vascular vs. Non-Vascular The reproductive adaptations of

angiosperms include flowers and fruits
Seed Plants Seedless Bryophytes  Angiosperms are flowering plants
 These seed plants have reproductive structures called flowers and fruits
Lycophyta (club/spike moss, Liverworts  They are the most widespread and diverse of all plants
quillworts) Hornworts
Pterophyta (ferns, horsetails, Mosses
whisk ferns)

Gymnosperms Angiosperms
(naked seeds) (container/fruit seeds)

Coniferophyta (Conifers)
Monocots Dicots
Cyadophyta (Cyads—palm-like)
Ginkophyta (Ginkos)
Gnetophyta

51 52

Characteristics of Angiosperms Flowers

 All angiosperms are classified in a single phylum, Anthophyta
 The name comes from the Greek anthos, flower
 The flower is an angiosperm structure specialized for sexual reproduction
 A flower is a specialized shoot with up to four types of modified leaves:
 Sepals, which enclose the flower
 Petals, which are brightly colored and attract pollinators
 Stamens, which produce pollen
 Carpels, which produce ovules

53 54

9
 LE 30-7
Fruits
Stigma
Carpel
Stamen Anther
Style
Filament  A fruit typically consists of a mature ovary but can also include other
Ovary
flower parts
 Fruits protect seeds and aid in their dispersal

 M ature fruits can be either fleshy or dry

Petal

Sepal

Ovule
Receptacle

55 56

Also, cereals,
Ruby grapefruit, a fleshy fruit
wheat, etc. Tomato, a fleshy fruit with soft with a hard outer layer and soft
in grasses outer and inner layers of pericarp inner layer of pericarp

 Various fruit adaptations help disperse seeds

 Seeds can be carried by wind, water, or animals to new locations

Nectarine, a fleshy
fruit with a soft outer
layer and hard inner
layer (pit) of pericarp

Milkweed, a dry fruit that splits Walnut, a dry fruit that remains
open at maturity closed at maturity

57 58

LE 30-9
Wings enable maple
fruits to be easily
Angiosperm Evolution
carried by the wind.

 Clarifying the origin and diversification of angiosperms poses fascinating

challenges to evolutionary biologists
 Angiosperms originated at least 140 million years ago
 During the late M esozoic, the major branches of the clade diverged
from their common ancestor
Seeds within berries
and other edible fruits
are often dispersed in
animal feces.

The barbs of cockleburs

facilitate seed dispersal
by allowing these fruits to
hitchhike on animals.

59 60

10
 LE 30-11
Fossil Angiosperms Carpel

Stamen

 Primitive fossils of 125-million-year-old angiosperms display derived and

primitive traits
 Archaefructus sinensis, for example, has anthers and seeds but lacks
petals and sepals

5 cm
Archaefructus sinensis, a
125-million-year-old fossil

Artist’s reconstruction of
Archaefructus sinensis

61 62

An “Evo-Devo” Hypothesis of Flower Angiosperm Diversity

Origins
 Scientist M ichael Frohlich hypothesized how pollen-producing and  The two main groups of angiosperms are monocots and eudicots
ovule-producing structures were combined into a single flower
 He proposed that the ancestor of angiosperms had separate pollen-
producing and ovule-producing structures

63 64

LE 30-12ba LE 30-12bb

MONOCOTS EUDICOTS

Pyrenean oak
MONOCOTS EUDICOTS Leaf (Quercus
venation pyrenaica)
Orchid California
Monocot Eudicot
(Lemboglossum poppy
rossii) Characteristics Characteristics
(Eschscholzia
california) Veins usually
Veins usually
Embryos parallel netlike

Stems
One cotyledon Two cotyledons
Pygmy date palm (Phoenix roebelenii)

Vascular tissue
Vascular tissue usually arranged
scattered in ring

65 66

11
 LE 30-12bc LE 30-12bd

MONOCOTS EUDICOTS

MONOCOTS EUDICOTS Barley (Hordeum vulgare), a grass Pea

(Lathyrusner
vosus, Lord
Lily (Lilium Pollen Anson’s
“Enchantment”) blue pea),
a legume

Pollen grain with Pollen grain with

one opening three openings
Roots

Flowers
Dog rose (Rosa canina), a wild rose
Root system Taproot (main root) Anther Zucchini
usually fibrous usually present (Cucurbita
(no main root) Stigma Floral organs Floral organs usually Pepo), female
usually in in multiples of (left), and
Ovary multiples of three four or five male flowers
Filament

67 68

LE 30-13
Evolutionary Links Between Angiosperms and Animals

 Pollination of flowers by animals and transport of seeds by animals are

two important relationships in terrestrial ecosystems

A flower pollinated by A flower pollinated by A flower pollinated by nocturnal animals.

honeybees. hummingbirds.

69 70

Concept 30.4: Human welfare depends Products from Seed Plants

greatly on seed plants
 No group of plants is more important to human survival than seed plants  M ost of our food comes from angiosperms
 Plants are key sources of food, fuel, wood products, and medicine  Six crops (wheat, rice, maize, potatoes, cassava, and sweet potatoes)
yield 80% of the calories consumed by humans
 Our reliance on seed plants makes preservation of plant diversity critical
 M odern crops are products of relatively recent genetic change
resulting from artificial selection
 M any seed plants provide wood

 Secondary compounds of seed plants are used in medicines

71 72

12
 Threats to Plant Diversity

 Destruction of habitat is causing extinction of many plant species

 Loss of plant habitat is often accompanied by loss of the animal species
that plants support

73 74

Other Threats to Plant Diversity Introduced kudzu thriving in

South Carolina
 Invasive plants (i.e., oriental bittersweet, kudzu,
phragmites)
(http://www.invasiveplants.net/)
 Loss of pollinators:
colony collapse disorder
(http://www.ars.usda.gov/News/docs.htm?docid=
15572)
Loss of pollinating bats?*
(http://www.nytimes.com/2008/03/25/science/25b
ats.html?em&ex=1206590400&en=659fa34c5520a3
6c&ei=5087%0A)
*These bats are primarily insectivorous bats, such as little brown, Indiana and
small-footed bats, but condition may spread.

75 76

Oriental Bittersweet
 Oriental bittersweet is a vigorously
Colony Collapse Disorder
 Bee pollination is responsible for $15 billion in added crop value,
growing vine that climbs over and particularly for specialty crops such as almonds and other nuts, berries,
smothers vegetation which may die fruits, and vegetables.
from excessive shading or breakage.
 About one mouthful in three in the diet directly or indirectly benefits
 Oriental bittersweet currently occurs in a number of states from New York to from honey bee pollination.
North Carolina, and westward to Illinois.
 Beginning in October 2006, some beekeepers began reporting losses of
 When bittersweet climbs high up on trees the increased weight can lead to 30-90 percent of their hives.
uprooting and blow-over during high winds and heavy snowfalls. In addition,
Oriental bittersweet is displacing our native American bittersweet (Celastrus  The main symptom of CCD is simply no or a low number of adult honey
scandens) through competition and hybridization. bees present but with a live queen and no dead honey bees in the
hive.

77 78

13

Why are bees disappearing?
There are three major possibilities that are being looked into by
researchers:
 Pesticides may be having unexpected negative effects on
honey bees.
 A new parasite or pathogen may be attacking honey bees. One
possible candidate being looked at is a pathogenic gut microbe
called Nosema. Viruses are also suspected.
 A perfect storm of existing stresses may have unexpectedly
weakened colonies leading to collapse. Stress, in general,
compromises the immune system of bees (and other social
insects) and may disrupt their social system, making colonies
more susceptible to disease.
 These stresses could include high levels of infection by the varroa
mite (a parasite that feeds on bee blood and transmits bee
viruses); poor nutrition due to apiary overcrowding, pollination of
crops with low nutritional value, or pollen or nectar scarcity; and
exposure to limited or contaminated water supplies. Migratory
stress brought about by increased needs for pollination might
also be a contributing factor.
79
Israeli acute paralysis virus (IAPV)
 The only pathogen found in almost all samples from
honey bee colonies with CCD, but not in non-CCD
colonies, was the Israeli acute paralysis virus (IAPV), a
dicistrovirus that can be transmitted by the varroa
mite. It was found in 96.1 percent of the CCD-bee
samples.
 This research found was strictly a strong correlation of
the appearance of IAPV and CCD together—it does
not identify IAPV as the cause of CCD. No cause-and-
effect connection can be inferred from the genetic
screening data.
 IAPV was initially identified in honey bee colonies in
Israel in 2002, where the honey bees exhibited unusual
behavior, such as twitching wings outside the hive and
a loss of worker bee populations.
 The study also found IAPV in honey bees from Australia
that had been imported into the United States, as well
as in royal jelly imported from China.
80
14