17.

The Evolution
of Sex

In the previous section, we mentioned some of the structural differences between prokaryotes and eukaryotes.
But one of the most profoundly important characteristics
of eukaryotes is the capacity for sexual reproduction. Indeed,
many types of protists undergo sexual reproduction. In sexual
reproduction, two different parents contribute gametes to
form the offspring. Gametes are usually formed by meiosis,
discussed in chapter 9. In most eukaryotes, the gametes are
haploid (have a single copy of each chromosome), and the
offspring produced by their fusion are diploid (have two copies of each chromosome). In this section, we examine sexual
reproduction among the eukaryotes and how it evolved.

Paramecium cells

(a)

Nuclei

Life Without Sex

To fully understand sexual reproduction, we must first examine asexual reproduction among the eukaryotes. Consider, for
example, a sponge. A sponge can reproduce by simply fragmenting its body, a process called budding. Each small portion
grows and gives rise to a new sponge. This is an example of
asexual reproduction, reproduction without forming gametes.
In asexual reproduction, the offspring are genetically identical to the parent, barring mutation. The majority of protists
reproduce asexually most of the time. Some protists such as
the green algae exhibit a true sexual cycle, but only transiently.
Asexual reproduction in a protist called Paramecium is shown
in figure 17.3a. The single cell duplicates its DNA, grows
larger, and then splits in two. The fusion of two haploid cells
to create a diploid zygote, the essential act of sexual reproduction, occurs only under stress. Paramecium is again shown in
figure 17.3b but now undergoing sexual reproduction. In this
case, the cell is not splitting in half; rather two cells are coming into close contact. In a process called conjugation, they
exchange genetic information in their haploid nuclei.
The development of an adult from an unfertilized egg
is a form of asexual reproduction called parthenogenesis.
Parthenogenesis is a common form of reproduction among
insects. In bees, for example, fertilized eggs develop into females, while unfertilized eggs become males. Some lizards,
fishes, and amphibians reproduce by parthenogenesis; an unfertilized egg undergoes mitosis without cytokinesis to produce a diploid cell, which then undergoes development as if it
had been produced by sexual union of two gametes.
Many plants and marine fishes undergo a form of
sexual reproduction that does not involve partners. In selffertilization, one individual provides both male and female
gametes. Mendels peas, discussed in chapter 10, produced
their F2 generations by selfing. Why isnt this asexual reproduction (after all, there is only one parent)? This is considered to be sexual rather than asexual reproduction because
the offspring are not genetically identical to the parent. During the production of the gametes by meiosis, considerable
genetic reassortment occursthat is why Mendels F2 plants
were not all the same!
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Paramecium
cells

(b)

Figure 17.3 Reproduction among paramecia.

(a) When Paramecium reproduces asexually, a mature individual
divides, and two genetically identical individuals result. (b) In sexual
reproduction, two mature cells fuse in a process called conjugation
(100) and exchange haploid nuclei.

Why Sex?
If reproduction without sex is so common among eukaryotes
today, it is a fair question to ask why sex occurs at all. Evolution is the result of changes that occur at the level of individual survival and reproduction, and it is not immediately
obvious what advantage is gained by the progeny of an individual that engages in sexual reproduction. Indeed, the segregation of chromosomes that occurs in meiosis tends to disrupt advantageous combinations of genes more often than it
assembles new, better-adapted ones. Because all the progeny
could maintain a parents successful gene combinations if the
parent employed asexual reproduction, the widespread use of
sexual reproduction among eukaryotes raises a puzzle: Where
is the benefit from sex that promoted the evolution of sexual
reproduction?

How Sex Evolved

In attempting to answer this question, biologists have looked
more carefully at where sex first evolvedamong the protists. Why do many protists form a diploid cell in response
to stress? Biologists think this occurs because only in a diploid cell can certain kinds of chromosome damage be repaired effectively, particularly double-strand breaks in DNA.
Such breaks are induced, for example, by desiccationdrying
out. The early stages of meiosis, in which the two copies of
each chromosome line up and pair with each other, seems to

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Key:

Diploid

Haploid

Haploid
cells

+
+

Gametes
+
+

Meiosis

2n
Reproductive
cell
2n

2n
Zygote

Haploid
individuals

+
Gametes

+
Gametes

Spores
+
+

Meiosis

Zygote

Diploid
individual

2n
Syngamy

(a) Zygotic meiosis

(b) Gametic meiosis

2n

Spore-forming
cell

2n

Gametophytes
(haploid)
n

+
Gametes

Syngamy

Meiosis

Sporophyte
(diploid)
2n Zygote

Syngamy

(c) Sporic meiosis

Figure 17.4 Three types of eukaryotic life cycles.

(a) Zygotic meiosis, a life cycle found in most protists. (b) Gametic meiosis, a life cycle typical of animals. (c) Sporic meiosis, a life cycle found
in plants.

have evolved originally as a mechanism for repairing doublestrand damage to DNA by using the undamaged version of the
chromosome as a template to guide the fixing of the damaged
one. In yeasts, mutations that inactivate the system that repairs
double-strand breaks of the chromosomes also prevent crossing over. Thus, it seems likely that sexual reproduction and the
close association between pairs of chromosomes that occurs
during meiosis first evolved as mechanisms to repair chromosomal damage by using the second copy of the chromosome
as a template.

Why Sex Is Important

One of the most important evolutionary innovations of eukaryotes was the invention of sex. Sexual reproduction provides
a powerful means of shuffling genes, quickly generating different combinations of genes among individuals. Genetic
diversity is the raw material for evolution. In many cases
the pace of evolution appears to be geared to the level of
genetic variation available for selection to act uponthe
greater the genetic diversity, the more rapid the evolutionary pace. Programs for selecting larger domestic cattle and
sheep, for example, proceed rapidly at first but then slow
as all of the existing genetic combinations are exhausted;
further progress must then await the generation of new gene
combinations. The genetic recombination produced by sexual reproduction has had an enormous evolutionary impact
because of its ability to rapidly generate extensive genetic
diversity.

Sexual Life Cycles

Many protists are haploid all their lives, but with few exceptions, animals and plants are diploid at some stage of their
lives. That is, the body cells of most animals and plants have
two sets of chromosomes, one from the male and one from the
female parent. The production of haploid gametes by meiosis,
followed by the union of two gametes in sexual reproduction,
is called the sexual life cycle.

Eukaryotes are characterized by three major types of

sexual life cycles (figure 17.4):
1. In the simplest of these, found in many algae, the
zygote formed by the fusion of gametes is the only
diploid cell. This sort of life cycle, which you can see
in figure 17.4a, is said to represent zygotic meiosis,
because in algae the zygote undergoes meiosis. Haploid
cells occupy the major portion of the life cycle, as
indicated by the larger yellow box; the diploid zygote
undergoes meiosis immediately after it is formed.
2. In most animals, the gametes are the only haploid cells.
They exhibit gametic meiosis, because in animals
meiosis produces the gametes. Here the diploid cells
occupy the major portion of the life cycle, as indicated
by the larger blue box in figure 17.4b.
3. Plants exhibit sporic meiosis, because in plants the
spore-forming cells undergo meiosis. In plants there is
a regular alternation of generations between a haploid
phase (the yellow boxed area in figure 17.4c) and a
diploid phase (the blue boxed area in figure 17.4c).
The diploid phase produces spores that give rise to the
haploid phase, and the haploid phase produces gametes
that fuse to give rise to the diploid phase.
The genesis of sex, then, involved meiosis and fertilization
with the participation of two parents. We have previously said
that bacteria lack true sexual reproduction, although in some
groups, two bacteria do pair up in conjugation and exchange
parts of their genome. The evolution of true sexual reproduction
among the protists has no doubt contributed importantly to their
tremendous diversification and adaptation to an extraordinary
range of ways of life, as we shall see in section 17.3.
Key Learning Outcome 17.2 Sex evolved among
eukaryotes as a mechanism to repair chromosomal
damage, but its importance is as a means of
generating diversity.

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The Protists

17.3

General Biology
of Protists, the
Most Ancient
Eukaryotes

Protists are the most ancient eukaryotes and are united on the
basis of a single negative characteristic: They are not fungi,
plants, or animals. In all other respects, they are highly variable with no uniting features. Many are unicellular, like the
Vorticella you see in figure 17.5 with its contractible stalk, but
there are numerous colonial and multicellular groups. Most
are microscopic, but some are as large as trees. We will start
our discussion of the protists with an overview of some of
their important features.

The Cell Surface

Protists possess varied types of cell surfaces. All protists have
plasma membranes. But some protists, like algae and molds,
are additionally encased within strong cell walls. Still others,
like diatoms and radiolarians, secrete glassy shells of silica.

Main body of
Vorticella cell

Cilia

Contractile
stalk

Substrate to
which this
Vorticella is
attached

Locomotor Organelles
Movement in protists is also accomplished by diverse mechanisms. Protists move by cilia, flagella, pseudopods, or gliding
mechanisms. Many protists wave one or more flagella to propel themselves through the water, whereas others use banks of
short, flagella-like structures called cilia to create water currents for their feeding or propulsion. Pseudopodia are the chief
means of locomotion among amoebas, whose pseudopods are
large, blunt extensions of the cell body called lobopodia. Other
related protists extend thin, branching protrusions called filopodia. Still other protists extend long, thin pseudopodia called
axopodia supported by axial rods of microtubules. Axopodia
can be extended or retracted. Because the tips can adhere to
adjacent surfaces, the cell can move by a rolling motion, shortening the axopodia in front and extending those in the rear.

Cyst Formation
Many protists with delicate surfaces are successful in quite
harsh habitats. How do they manage to survive so well? They
survive inhospitable conditions by forming cysts. A cyst is
a dormant form of a cell with a resistant outer covering in
which cell metabolism is more or less completely shut down.
Amoebic parasites in vertebrates, for example, form cysts that
are quite resistant to gastric acidity (although they will not
tolerate desiccation or high temperature).

Nutrition
Protists employ every form of nutritional acquisition except
chemoautotrophy, which has so far been observed only in
prokaryotes. Some protists are photosynthetic autotrophs and
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Figure 17.5 A unicellular protist.

The protist kingdom is a catch-all kingdom for many different groups
of unicellular organisms, such as this Vorticella (phylum Ciliophora),
which is heterotrophic, feeds on bacteria, and has a contractible stalk.

are called phototrophs. Others are heterotrophs that obtain

energy from organic molecules synthesized by other organisms. Among heterotrophic protists, those that ingest visible
particles of food are called phagotrophs, or holozoic feeders.
Those ingesting food in soluble form are called osmotrophs,
or saprozoic feeders.
Phagotrophs ingest food particles into intracellular vesicles called food vacuoles, or phagosomes. Lysosomes fuse
with the food vacuoles, introducing enzymes that digest the
food particles within. As the digested molecules are absorbed
across the vacuolar membrane, the food vacuole becomes progressively smaller.

Reproduction
Protists typically reproduce asexually, most reproducing sexually only in times of stress. Asexual reproduction involves
mitosis, but the process is often somewhat different from the
mitosis that occurs in multicellular animals. The nuclear membrane, for example, often persists throughout mitosis, with
the microtubular spindle forming within it. In some groups,
asexual reproduction involves spore formation, in others fission. The most common type of fission is binary, in which a
cell simply splits into nearly equal halves. When the progeny

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cell is considerably smaller than its parent, and then grows to

adult size, the fission is called budding. In multiple fission,
or schizogony, common among some protists, fission is preceded by several nuclear divisions, so that fission produces
several individuals almost simultaneously.
Sexual reproduction also takes place in many forms
among the protists. In ciliates, gametic meiosis occurs just
before gamete formation, as it does in most animals. In the
sporozoans, zygotic meiosis occurs directly after fertilization, and all the individuals that are produced are haploid until
the next zygote is formed. In algae, there is sporic meiosis,
producing an alternation of generations similar to that seen
in plants, with significant portions of the life cycle spent as
haploid as well as diploid.

Multicellularity
A single cell has limits. It can only be so big without encountering serious surface-to-volume problems. Said simply, as a
cell becomes larger, there is too little surface area for so much
volume. The evolution of multicellular individuals composed
of many cells solved this problem. Multicellularity is a condition in which an organism is composed of many cells, permanently associated with one another, that integrate their activities. The key advantage of multicellularity is that it allows
specializationdistinct types of cells, tissues, and organs
can be differentiated within an individuals body, each with
a different function. With such functional division of labor
within its body, a multicellular organism can possess cells devoted specifically to protecting the body, others to moving it
about, still others to seeking mates and prey, and yet others to
carry on a host of other activities. This allows the organism
to function on a scale and with a complexity that would have
been impossible for its unicellular ancestors. In just this way,
a small city of 50,000 inhabitants is vastly more complex and
capable than a crowd of 50,000 people in a football stadium
each city dweller is specialized in a particular activity that is
interrelated to everyone elses, rather than just being another
body in a crowd.

given area and are near starvation, all of the individual

organisms in that immediate area aggregate into a large
moving mass of cells called a slug. By moving to a different location, the aggregation increases the chance that
food will be found.
Multicellular Individuals True multicellularity, in which
the activities of the individual cells are coordinated and
the cells themselves are in contact, occurs only in eukaryotes and is one of their major characteristics. Three groups
of protists have independently attained true but simple
multicellularitythe brown algae (phylum Phaeophyta),
green algae (phylum Chlorophyta), and red algae (phylum
Rhodophyta). In multicellular organisms, individuals are
composed of many cells that interact with one another and
coordinate their activities.
Simple multicellularity does not imply small size or
limited adaptability. Some marine algae grow to be enormous. An individual kelp, one of the brown algae, may
grow to tens of meters in lengthsome taller than a redwood! Red algae grow at great depths in the sea, far below where kelp or other algae are found. But not all algae
are multicellular. Green algae, for example, include many
kinds of multicellular organisms but an even larger number of unicellular ones.
Key Learning Outcome 17.3 Protists exhibit
a wide range of forms, locomotion, nutrition,
and reproduction. Their cells form clusters with
varying degrees of specialization, from transient
aggregations to more persistent colonies to
permanently multicellular organisms.

Colonies A colonial organism is a collection of cells that are

permanently associated but in which little or no integration of
cell activities occurs. Many protists form colonial assemblies,
consisting of many cells with little differentiation or integration.
In some protists, the distinction between colonial and multicellular is blurred. For example, in the green algae Volvox shown
in figure 17.6, individual motile cells aggregate into a hollow
ball of cells that moves by a coordinated beating of the flagella
of the individual cellslike scores of rowers all pulling their
oars in concert. A few cells near the rear of the moving colony
are reproductive cells, but most are relatively undifferentiated.
Aggregates An aggregation is a more transient collection
of cells that come together for a period of time and then
separate. Cellular slime molds, for example, are unicellular organisms that spend most of their lives moving about
and feeding as single-celled amoebas. They are common
in damp soil and on rotting logs, where they move around
and ingest bacteria and other small organisms. When the
individual amoebas exhaust the supply of bacteria in a

Figure 17.6 A colonial protist.

Individual, motile, unicellular green algae are united in the protist
Volvox as a hollow colony of cells that moves by the beating of
the flagella of its individual cells. Some species of Volvox have
cytoplasmic connections between the cells that help coordinate
colony activities. The Volvox colony is a highly complex form that
has many of the properties of multicellular life.
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