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BIO 152 LAB 10 Animal Developemnt Worksheet

Foundations of Biological Sciences II (University of Wisconsin-Milwaukee)

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BIO SCI 152 LABORATORY 10 – SPRING 2020 WORKSHEET

ANIMAL DEVELOPMENT I: ECHINODERMS & AMPHIBIANS

OBJECTIVES OF LABORATORY:
1. Compare and contrast early development in an echinoderm (sea star) and amphibian
(frog) identifying events that are in common and those that differ.
2. Understand the effects of egg yolk size and location on subsequent early development.
3. Relate the events of early development in vertebrates to the formation of a dorsal nerve
chord.
4. Understand the importance and mode of regeneration in some animals.

I. PATTERNS OF ANIMAL DEVELOPMENT – AN OVERVIEW

Animals come in a wide variety of shapes, sizes, and structure, ranging from mammals
(including humans) to insects, fish, earthworms, tapeworms, jellyfish, and sponges. Currently,
about 35 phyla of animals are recognized, but there are many challenges in understanding the
evolutionary relationships among taxa.

Embryology is important in understanding the evolution of body plans in animals. All of the
organs and systems of animals develop from a single cell, the zygote. Any subsequent changes in
organs and systems depend on changes in patterns of development. In this lab, we will explore
some of the basic features of animal embryology, and introduce some ways that changes in body
plan depend on simple differences in patterns of early development. Development of a
multicellular organism is the process by which a zygote is transformed into an adult organism,
including the production and fusion of male and female gametes, development of a multicellular
embryo, and emergence of larval or juvenile stages, growth and maturation to sexual maturity,
and eventually death. A range of biological processes function in development, including cell
division, differentiation (cells tissues and organs become specialized for a particular function),
and morphogenesis (the development of the animal’s body form and organization).

Developmental biologists began by studying morphology (form or shape of an organism) and by

describing the process of development. We now have a reasonably detailed understanding of the
genetic basis for development and the processes involved in activating different genes in
different cells. DNA dictates, through time, the structural and functional development of a single
cell into an embryo and its morphogenesis into an adult. Even aging is a result of genetic
programming! Recent molecular techniques have made it possible to map the entire genomes of
some organisms and find common patterns of gene function in animal development.

In this lab, you will investigate the major morphological events that take place in organisms
during early development. These events include; gametogenesis (the production of gametes),
fertilization, cleavage (rapid mitotic division without cell growth), and blastulation (the
production of a multicellular blastula), gastrulation (the formation of three primary germ layers
- ectoderm, mesoderm and endoderm), neurulation (formation of nervous system in chordates),
and organogenesis (development of organs from the three primary germ layers).

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II. OVERVIEW OF THE FOUR STAGES IN EARLY DEVELOPMENT

Stage 1: Preparation of Egg, Fertilization, and Cleavage
Development begins with the preparation of sperm and egg for fertilization. A sperm cell
develops a flagellum, used to propel the haploid gamete containing the paternal genome toward
the haploid egg containing the maternal genome. The egg contains food reserves of yolk, rich in
protein and fat that can be utilized by the early embryo. When egg and sperm come into contact,
their nuclei combine to form a diploid zygote. Subsequent mitotic divisions, called cleavage,
rapidly convert the zygote to a multicellular ball or disc called a blastula. The cells of the
blastula are called blastomeres (Gr: blastus = a bud, mere = part). Eventually, a cavity, the
blastocoel, forms within the ball of cells. The location of the blastocoel varies according to the
egg type.

Egg types. Eggs are classified according to the amount and distribution of yolk in the egg, which
affects subsequent early development. Isolecithal (“iso” means same and lecithal refers to yolk)
eggs contain small amounts of evenly distributed yolk (Figure 1). Isolecithal eggs are
characteristic of echinoderms, molluscs, and mammals. Telolecithal (“telo” means end) eggs
contain large amounts of yolk concentrated at one end. Some amphibian species are
mesolecithal, while reptiles, birds, and fish are telolecithal.

In telolecithal eggs, there are two hemispheres: the animal hemisphere, which contains small
cells that divide rapidly and the vegetal hemisphere which has large yolk cells that divide
slowly (Figure 1). The animal pole (in the animal hemisphere) contains the nucleus and is
surrounded by active cytoplasm that is relatively devoid of yolk (you may have noticed this
region in eggs when cooking.) This nuclear-cytoplasm region is called the blastodisc. The
blastodisc is toward the pole of the egg where polar bodies budded from the cell during meiosis.
In these eggs, the yolk is concentrated in the other half of the egg (vegetal hemisphere), near the
vegetal pole. In frogs, a single sperm can enter anywhere on the animal hemisphere of the egg;
when it does, it changes the cytoplasmic pattern of the egg. Originally, the egg is radially
symmetrical about the animal-vegetal axis. After sperm entry, however, the cortical (outer)
cytoplasm shifts about 30° toward the point of sperm entry, relative to the inner cytoplasm. A
region of the egg that was formerly covered by the dark cortical cytoplasm of the animal
hemisphere is now and appears gray. This region is referred to as the gray crescent and indicates
the region where gastrulation is initiated in amphibian embryos.

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Figure 1. Reorganization of cytoplasm in the newly fertilized frog egg.

Cleavage types. The end result of cleavage, the formation of the blastula, is the same in all
multicellular animals. However, the pattern of cleavage can differ among organisms depending
on egg type. In isolecithal eggs, where the impact of the yolk is minimal, holoblastic cleavage
occurs with the cell divisions passing through the entire fertilized egg. In these eggs, the
blastocoel forms in the center of the blastula (Figure 2). In mesolecithal eggs, the yolk will
retard cytoplasmic divisions and affect the size of the cells. If the entire egg is cleaved, as
demonstrated in isolecithal and mesolecithal eggs, cleavage is considered holoblastic. In this case
the blastocoel develops in the animal hemisphere. Cells in this hemisphere will be smaller and
have less yolk than cells in the vegetal hemisphere. In telolecithal eggs, only the active
cytoplasm is divided during cleavage. This is known as meroblastic cleavage and it produces a
cap of cells known as a blastoderm. The blastocoel forms between two layers of cells within the
blastoderm.
Isolecithal Egg
Radial cleavage Holoblastic = complete cleavage
through entire cell

Spiral cleavage Cleavage begins at the “animal”

pole

Bilateral cleavage

Rotational cleavage

Mesolecithal Egg Cleavage can be delayed yolk;

Radial cleavage larger cells in vegetal pole

Telolecithal Egg Meroblastic = incomplete cleavage

Discoidal cleavage

Figure 2. Effect of yolk content on cleavage patterns.

Stage 2: Gastrulation
Gastrulation transforms the blastula into a gastrula made up of three embryonic (germ) layers:
endoderm, ectoderm and mesoderm (Figure 3). Whereas cleavage is characterized by cell
division, gastrulation is characterized by cell movement. Surface cells migrate into the interior of
the embryo in a process called involution, forming a new internal cavity, the archenteron. The
archenteron is lined by the endoderm, the embryonic germ layer that will eventually form the
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digestive tract. The archenteron opens to the outside through the blastopore. The cells on the
surface of the embryo become the ectoderm. The third layer of cells, the mesoderm, develops
between the ectoderm and endoderm. The future function of the blastopore is an important
characteristic that defines two major lineages within the animals: the protostomes and the
deuterostomes. In protostomes, the blastopore becomes the mouth. In deuterostomes, the
blastopore becomes the anus.

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Figure 3. Gastrulation and the development of the 3 primary embryonic layers.

Stage 3: Neurulation
Late in gastrulation, neurulation begins. Neurulation is the formation of a dorsal hollow neural
tube, found only in the chordates. Certain ectodermal cells flatten into an elongated neural
plate, extending from the dorsal edge of the blastopore to the anterior end of the embryo. The
center of the plate sinks, forming the neural groove (Figure 4B). The edges of the plate become
elevated to form neural folds, which gradually approach each other and eventually fuse, forming
the hollow neural tube. The anterior end of the neural tube develops into the brain; the posterior
end develops into the nerve or spinal cord.

Figure 4. Neurulation and the formation of the neural

tube.
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Stage 4: Organogenesis
After the germ layers and the nervous system have been established, organogenesis, the
formation of rudimentary organs and organ systems, begins. The ectoderm will develop into
skin and associated glands. In chordates, somites and the notochord develop early from
mesodermal cells. Somites form in pairs flanking the neural tube (Figure 4D). Somites are
blocks of cells that form a segmental pattern, eventually developing into vertebrae, ribs,
muscles, and skin (Figure 5). The skeleton, gonads, the excretory system, and the circulatory
system also develop from the mesoderm. Non-chordate animals lack somites and a notochord,
but their muscles and organs of excretory, circulatory and reproductive systems develop from
the mesoderm. The endoderm develops into the lining of the digestive tract and associated
organs, such as the liver, pancreas, and lungs.

Somites

Figure 5. Somites in a two-day chicken embryo

III.DEVELOPMENT OF AN ECHINODERM: THE SEA STAR

Sea stars belong to the Phylum Echinodermata, an invertebrate group that is phylogenetically
closer to the chordate lineage than any other group. They display the typical deuterostome
pattern of early development. Male and female sea stars release large numbers of gametes into
the sea, and fertilization is external. Early development leads to a larval stage that is free-
swimming and free- feeding.

In this exercise, you will observe a slide that contains an assortment of whole embryos in various
stages of development. Identify each developmental stage and determine the type of egg and
cleavage pattern found in echinoderms. Make careful drawings in the spaces provided below.

Unfertilized egg
By the time sea star eggs leave the body of the female, meiosis I and II have been completed.
The nucleus, called the germinal vesicle, is conspicuous because the nuclear envelope is intact. A
nucleolus is usually distinct. The plasma membrane surrounding the egg cytoplasm closely
adheres to a thin external membrane known as the vitelline layer that contains species-specific
sperm receptors.
1) Draw the unfertilized egg below. Label nucleus, nucleolus, and vitelline layer, if visible.

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Fertilized egg
The fertilized egg, or zygote, has no visible nuclear envelope, giving the cell a uniform
appearance. The zygote surface develops a fertilization membrane. This membrane forms after
sperm—egg fusion and helps prevent polyspermy (multiple fertilizations). Two sequential
processes prevent polyspermy, the fast block and the slow block (Figure 6).

Figure 6. Sequence of events upon initial contact of the sperm with a sea urchin egg.

2) Draw a fertilized egg below. Label the fertilization membrane and other evident features.

Early Cleavage
The earliest stages of cleavage are named for the number of cells they contain: the 2-cell stage,
the 4-cell stage, the 8-cell stage, and so on. Individual cells that form during these cleavages are

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called blastomeres.

3) Find and draw below the 2-, 4-, and 8-cell stages on your slide.

4) Is the entire zygote involved in early cleavage? What evidence to you have to support your
answer?
Yes, the entire zygote is involved in the cleavage. There are different patterns of
cleavage depending on the egg type when a single celled zygote changes into
multicellular blastula or gastrula.

5) What happens to the size of the cells as cleavage takes place?

The size of the cell decreases as cleavage decreases

Late Cleavage
Once the cells become too numerous to be counted easily, which usually happens around the 32-
or 64-cell stage, this terminology is abandoned and the embryo is said to be in the morula stage.
The morula is a spherical mass of from a few dozen to several hundred cells.

6) Locate the morula stage on your slide and draw it below.

Blastulation
The morula is solid, but as cleavage continues, the blastomeres move toward the periphery of the
mass, leaving a fluid-filled cavity in the interior. Soon the embryo no longer resembles a simple

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mass of cells, but instead takes on the form of a sphere. In this stage the embryo is called a
blastula, and the fluid-filled cavity is called a blastocoel (Gr: coelo = hollow). This change takes
place about ten hours after fertilization. When the blastula first forms it contains about 1000
cells.

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7) Locate and study a blastula on your slide and draw it below. Label the blastocoel if visible.

8) How does the size of individual cells compare with the size of the fertilized egg?

The individual cells are much smaller than the fertilized egg.

9) How does the overall size of the blastula compare with the size of the fertilized egg?

The blastula will be larger than the fertilized egg.

Early Gastrulation
A few hours after the formation of the blastula the process of gastrulation begins. Gastrulation
converts the blastula into a gastrula, an embryo composed of three germ layers. At a certain point on
the surface of the blastula, cells begin to move toward the center of the blastocoel, so that an
opening is formed leading into a blind or dead-end tube. If you were to take a rubber balloon loosely
filled with water or air and poke your finger into it without breaking the balloon you would be at
least roughly approximating the process of gastrulation.
The opening on the surface of the embryo is called the blastopore, and in deuterostomes such as sea
urchins and sea stars, it will become the anus of the mature organism. The blind or dead-end tube is
called the archenteron (Gr: archi = first, enterum = intestine). This will eventually reach all the way
through and open out the other side and will form the basis of the animal’s digestive tract. Once a
blastopore and an archenteron have formed, the embryo is called a gastrula (Gr: gastro = stomach).
The early gastrula can be recognized by a small swelling of cells protruding into the blastocoel.
10) Draw the early stage of gastrulation below.

11) Which embryonic germ layer lines the archenteron?

The endoderm lines the archenteron and gives rise to the liver, pancreas, lungs, and
the lining of the digestive tract.

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Middle Gastrulation
The archenteron continues to grow across the blastocoel. It takes on a bulb-like appearance as
the advancing portion swells.

12) Draw and label the gastrula, archenteron, and blastopore.

Late Gastrulation
At the gastrula stage of development a certain amount of differentiation has taken place. The
cells on the outside of the gastrula constitute the ectoderm (Gr: ecto = outer, derma = skin), and
these cells will develop into the epidermis and the nervous system of the mature animal. The
cells of the archenteron constitute the endoderm (Gr: endo = inside), and these will become the
lining of the digestive tract and the digestive glands of the fully developed organism. These two
primary germ layers will be joined by a third one, the mesoderm. The amoeboid cells that attach
the archenteron to the embryo wall are called mesenchyme cells. These cells later detach from
the archenteron, proliferate, and form a layer of cells lining the old blastocoel, now divided by
the archenteron. This layer of cells will become the mesodermal germ layer.

13) Draw the late gastrulation embryo below. Label the blastopore, archenteron, and mouth.

14) What will be formed from the blastopore?

During maturation of some animals it evolves into the anus or the mouth; in others it is
covered over and contributes to the canal joining the primitive gut with the cavity of
15) What is the germ layer of cells on the surface of the embryo called?

The ectoderm is the outside germ layer of the embryo.

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Bipinnaria Larval Stage

As sea urchins and sea stars grow, differentiation and morphogenesis continue beyond the
gastrula stage until a free-living bipinnaria larval stage is formed. This bilaterally symmetric
larvae is fully capable of moving about and feeding itself. The archenteron of the gastrula
differentiates into a broad esophagus leading from the mouth to a large oval stomach and on to a
small tubular intestine. All these structures should be visible on your slide. At a later stage, a
metamorphosis will occur and the larva will assume the form of the radially symmetric adult
starfish.

16) In the micrograph below of a larval sea urchin, complete the blanks, labeling the mouth,
blastopore (anus), archenteron, blastocoel, and stomach.

Blastocoel

Blastopore
Archenteron

Stomach

Mouth

Summary questions:

17) What is the advantage of species-specific sperm receptors in the vitelline layer of the
egg?
It ensures that fertilization will only occur with sperm from the same species.

18) What type of egg does the sea star have? What evidence do you have to support your
answer?
Sea stars have isolecithal eggs. The yolk is almost evenly distributed within the egg,
which undergoes uniform cleavage.
19) Describe the pattern of cleavage seen in the sea star and give the name for this type of
cleavage?
The whole yolk is cleaved, which is called holoblastic cleavage.

IV. DEVELOPMENT IN AMPHIBIANS

Amphibians (frogs and salamanders) are vertebrates that lay jelly-coated eggs in water or moist
areas on land. For most species, fertilization is external; the male deposits sperm over the eggs
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after they are released by the female. However, internal fertilization does occur in some
amphibian species and the young are born in an advanced stage of development. Early
development is similar in all species, involving fertilization, cleavage, gastrulation, neurulation,
and organogenesis. Amphibians are a convenient system for studying vertebrate development for
several reasons: They produce large oocytes (mature African clawed toad oocytes are 1 mm in
diameter, an order of magnitude larger than a mouse oocyte), they undergo external development
and can readily be observed and manipulated, and development is relatively rapid (they go from
fertilization through neurulation in approximately 18 hours at 22°C). In this exercise, you will
study amphibian early development by observing microscope slides, models, and a video of
Xenopus development.

Answer the following questions based on your study of amphibian development.

20) Review common terms used in embryology by completing the table below:

Term Definition
the portion of an ovum that contains the nucleus and less yolk,
Animal pole
opposite the vegetal pole.
half of an egg or embryo that contains less yolk and/or which divides more
Animal hemisphere
rapidly in comparison to the vegetal hemisphere.
The location where kinetochore microtubules pull the sister chromatids
equator
back and forth until they.
the lower, yolky portion of the egg; opposite the animal hemisphere
Vegetal hemisphere
the portion of an ovum opposite the animal pole, containing most
Vegetal pole
of the yolk and little cytoplasm.

21) Is the amphibian egg isolecithal, mesolecithal, or strongly telolecithal?

The amphibian egg is telolecithal

22) Describe the cleavage pattern in amphibians. Is it holoblastic or meroblastic?

The cleavage is meroblastic as it occurs at the top of the egg.

23) Are cleavages synchronous or irregular? Can you detect any particular pattern in the
cleavage?
They are irregular with no patterns
24) Where is the second cleavage plane in relation to the first?
The second cleavage is done down the middle.

25) Does the size of the embryo change as cleavage progresses?

Yes, the embryo becomes larger.
26) Describe gastrulation in amphibians, comparing the process with the sea star.
Amphibians experience polar cleavage where only part of the zygote divides. In
comparison, the sea star undergoes a full zygote division.

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27) List at least two major differences between early development in the amphibian and the sea
star and describe the factors that may be responsible for those differences.
-one vs. two regions of embryo due to different cleavage
-type of cleavage is different (meroblastic for amphibians, holoblastic for sea stars)
-

28) Label the following in Figure 7: egg, sperm, fertilized egg, blastula, gastrula, neurula,
ectoderm, mesoderm, endoderm, blastocoel, and blastopore.
Fertilized egg Blastul
Sperm

Blastocoel
Egg
Blastopore
Ectoderm Neurula
Nautrala
Mesoderm
Endoderm
Gastrula

Figure 7. Frog life cycle.

V. REGENERATION IN LUMBRICULUS VARIEGATUS

This lab is adapted from: Drewes, C. D. 1996. Heads or Tails? Patterns of segmental
regeneration in a freshwater oligochaete. Tested studies for laboratory teaching, 17: 23-35.

Lumbriculus variegatus (common name blackworms) are found throughout North America and
Europe. They prefer shallow habitats at the edges of ponds, lakes, or marshes where they feed on
decaying vegetation and microorganisms. Lumbriculus uses its head to forage in sediments and
debris, while its tail end, specialized for gas exchange, often projects upwards. Maximal body
size in field-collected blackworms is about 10 cm in length and 1.5 mm in diameter.

Sexually mature blackworms are hermaphrodites (possess both male and female sexual organs).
Although never documented, sexual reproduction in mature worms probably involves copulation
and sperm exchange, as seen in many earthworms. Reproduction under natural and laboratory
conditions is usually by asexual fragmentation, during which a worm spontaneously divides
into two or more body fragments. The capacity for asexual reproduction by fragmentation is
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matched by the ability to self-amputate in response to injury or other types of stimulation. One
stimulus which causes worms to readily self-amputate is body compression. A sudden
compression stimulus, which presumably simulates a predatory attack, induces a rapid and clean
division from the body.

Regeneration
Many early diverging animals with relatively simple body plans are capable of regeneration. This
is a remarkable process in which portions of animals reorganize to form missing parts, usually
with remarkable speed and precision. Most organisms with more complex body plans (e.g.,
mammals) are incapable of regeneration, even though they originally had all the necessary
instructions and machinery to develop the tissues during early embryonic development.
Scientists are interested in understanding the mechanisms of regeneration in the hope that they
will be able to induce regeneration in non-regenerative systems (for example, humans).

Embryonic development is an orderly process during which an organism’s cells differentiate and
its body gradually acquires adult-like characteristics. If the adult organism is bilaterally
symmetrical (as in many invertebrates and all vertebrates), then at some time during
embryogenesis a body plan is established along three different body axes: (1) an anterior-
posterior axis that establishes head and tail ends, (2) a dorsal-ventral axis that establishes front
and back sides, and (3) a left-right axis. During development, the fate of differentiating cells and
tissues will vary, depending on their exact position within these axes. Cells closest to the head
end, under the influence of chemical factors (called morphogens) and physical constraints, will
develop proper, head-like features. Cells in the middle will be similarly influenced to develop
features appropriate to a mid-body position, etc. This complex developmental process of
acquiring a characteristic body plan with position-specific features is referred to as pattern
formation.

In some invertebrates (such as worms, hydra, and planarian) the process of pattern formation
may occur during the regeneration of body parts. If a worm loses a part of its posterior end by
fragmentation, how does the worm “know” whether to regrow a new head or tail at the wound
site? How does it “know” how long the replacement part should be? In relatively large and
segmented organisms, such as annelid worms, the underlying mechanisms of development are
not completely understood. However, we can begin to understand these mechanisms if we
carefully observe the regeneration process following various surgical manipulations, attempting
to determine the “rules” and general patterns that govern the process.

When organisms regrow missing body parts, one of two general regeneration patterns can occur.
In one pattern, morphallaxis, the majority of the regenerated tissue comes from tissue already
present in the organism. Reorganization of cells can occur without cell division. A classic
example of an organism that regenerates using this mechanism is the hydra (Figure 8). When a
hydra is cut into two parts, two hydra will regenerate, each smaller than the original hydra.
During the regenerative process very little cell division takes place. Once regeneration is
completed, the two hydra can continue to grow via cell division and reach the size of their
original parent.

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Figure 8. Regeneration by morphallaxis in Hydra, a freshwater cnidarian.

Morphallaxis is often contrasted with epimorphosis, which is characterized by a greater degree

of cellular proliferation. Epimorphosis can be further subdivided into dedifferentiation-
independent and dedifferentiation-dependent subclasses. Planarian regenerate using a
dedifferentiation-independent mechanism in which preexisting stem cells proliferate and migrate
to the injured site in response to injury. These cells form a mass of proliferating cells that later
differentiate into the specialized cells that become the regenerated structure. Most tissue
regeneration in mammals belongs to the dedifferentiation-independent subclass. For example,
mammals can regenerate their muscle, bone, epithelia tissue, and some neurons by activating
preexisting stem cells (progenitor cells). Certain vertebrates, such as the salamanders, regenerate
lost body parts through dedifferentiation-dependent epimorphosis. In this case, differentiated
cells reverse the normal developmental process and once again become precursor or stem cells.
These dedifferentiated cells then proliferate and re-differentiate to form the regenerated structure
or organ.

Although cellular differentiation is active in both processes, in morphallaxis the majority of the
regeneration comes from reorganization or exchange, while in epimorphosis the majority of the
generation comes from cellular differentiation. In this exercise you will determine which general
pattern of regeneration (morphallaxis or epimorphosis) occurs in Lumbriculus. You will
systematically compare head and tail regeneration in amputated fragments that are approximately
the same length (Figure 9). You will also determine whether short and long fragments from the
same body region have differing capacities for head and tail regeneration, and whether there is a
minimal size requirement.

30) State a hypothesis for which pattern of regeneration (morphallaxis or epimorphosis) you think
may occur in Lumbriculus. Explain why you chose this hypothesis.

I believe morphallaxis occurs during segmental regeneration of Lumbriculus as it has

the capacity in its cells to reproduce by using existing cells without cellular division.
Considering most injuries in Lumbriculus are restorative instead of reparative,
morphallaxis is a better mechanism.

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31) State a prediction for how the appearance of Lumbriculus will differ after regeneration by
either morphallaxis or epimorphosis.
In morphallaxis, new cells grow by the reorganization or exchange of the existing
cells so a reparation like the growth of a new tail would occur, whereas in
epimorphosis, majority of new cells grow by cellular differentiation, therefore the
restoration of an organ would occur due to the growth of specific job cells.

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