Lecture 15:  Development, Coelom Formation

 


I.       Early development

A.    The newly fertilized zygote is much larger than a normal body cell

1.                  The zygote divides a number of times until the cells of what is now the embryo are of normal somatic cell size

a)                  This process is called cleavage

b)                  During this time the embryo does not grow in size

c)                  Each one of the cells of the embryo is called a blastomere

(1)               At the end of cleavage, the number of cells varies (depending on species) from 1000 to 700,000

B.     Patterns of cleavage

1.                  Cleavage patterns are affected by

a)                  Quantity & distribution of yolk (Fig. 8-7)

(1)               Yolk retards the speed of cleavage – this means that different regions of the embryo may divide at different rates
(a)               Isolecithal egg – little yolk
(b)               Mesolecithal egg – moderate amount of yolk

(i)                  Gives egg polarity

(a)               End containing yolk is called vegetal pole

(b)               Opposite end is called animal pole – mostly cytoplasm & little yolk

(c)                In isolecithal & mesolecithal eggs, cleavage is holoblastic – the cleavage furrow extends completely through the egg
(d)               Telolecithal – lots of yolk concentrated at vegetal end

(i)                  Cleavage is telolecithal eggs is meroblastic – cleavage furrows extend only part way through the egg

b)                  Genes controlling the symmetry of cleavage (Fig. 3-6, Lab Manual)

(1)               Radial cleavage – cleavage furrows are oriented parallel or perpendicular to animal-vegetal pole axis
(a)               Animals that undergo radial cleavage are called the deuterostomia
(b)               Includes echinoderms, hemichordates, & chordates
(2)               Spiral cleavage – cleavage furrows are oriented obliquely to animal vegetal pole – blastomeres end up lying in cleavage furrows
(a)               Animals that undergo spiral cleavage are called the protostomia
(b)               Found in annelids, nemerteans, tubellarian flatworms, molluscs (except cephlopods), & several other invertebrate phyla
(3)               Protostome vs. deuterostome distinction is important in other aspects of development

c)                  Some other groups like insects (protostomes) & mammals (deuterostomes) have unique cleavage patterns (described in text)

II.    Blastulation (Fig 1; similar to Fig. 3-9, Lab Manual)

A.    Following cleavage, the embryo is a mass of cells called a blastula

1.                  In many animals, the cells surround a fluid-filled cavity called a blastocoel

III. Gastrulation

A.    Gastrulation converts a spherical blastula into a complex structure with 3 layers

B.     Pattern of gastrulation varies tremendously depending on the amount of yolk

1.                  Simple in non-yolky embryos, complex in yolk-laden embryos

C.    We’ll use the relatively simple gastrulation of the frog as our example

1.                  Gastrulation begins with the invagination of cells into the interior of the blastula

2.                  The invaginating cells move into the blastocoel & reduce its size (Fig. 1g-i)

3.                  During invagination, a new cavity is formed, the archenteron

a)                  The archenteron will form the gut

4.                  The opening of the archenteron is called the blastopore

a)                  In protostomes, the blastopore becomes the mouth, the anus will form secondarily (Fig. 8‑9)

b)                  In deuterostomes, the blastopore becomes the anus, the mouth forms secondarily

5.                  Note that the blastocoel is being obliterated during gastrulation

a)                  In some animals, the blastocoel will persist – more on that later

6.                  The embryo now has 2 cell layers, the outside layer is called ectoderm & the cells lining the archenteron are called endoderm

a)                  Most animals will form a cell layer between the ectoderm and endoderm, the mesoderm (Fig. 1j)

(1)               Cnidaria & Ctenophora do not form mesoderm

IV.  Coelom formation (Fig. 9-12)

A.    Many animals form a cavity inside the body called a coelom

1.                  Animals that do not form a coelom are called acoelomate

a)                  Acoelomates include flatworms & nemerteans

b)                  The area between the body wall & the gut is filled by a network of cells a parenchyma (mesodermal origin)

B.     Animals with a body cavity are divided into two large groups depending on how the body cavity is formed, pseudocoelomates & coelomates

1.                  In pseudocoelomates, the blastocoel persists after gastrulation & forms the pseudocoelom

a)                  The pseudocoelom is a perfectly good body cavity, but is called a “pseudocoelom” because the cavity isn’t lined by a mesodermal peritoneum (gut isn’t lined [surrounded] by mesoderm)

b)                  Organs derived from the mesoderm are found within the pseudocoelom

c)                  Pseudocoelomates include:  nematodes, rotifers, gastrotrichs

2.                  Coelomates form a “true coelom” in one of 2 ways (Fig. 9-13)

a)                  Protostomes form their coelom via a process called schizocoelous development

(1)               Mesoderm forms from cells near lips of blastopore
(2)               Mesoderm splits (schizo = split) to form coelom

b)                  Most deuterostomes form their coelom via enterocoelous development

(1)               Mesoderm forms from outpocketings of the archenteron

c)                  In both schizocoelous & enterocoelous development the coelom enlarges to obliterate the blastocoel

(1)               The mesodermal peritoneum will now line both the body wall & gut, with a double layer mesentery between the body wall & gut
(a)               The mesoepithelium you viewed in lab was this mesentery
(2)               Note that the organs do not lie within the coelom itself
(a)               All mesodermal organs are covered by the peritoneum (simple squamous epithelium)

d)                  Vertebrates are deuterostomes that do not form the coelom via enterocoelous development

(1)               Vertebrates produce their coelom via a secondarily derived schizocoelous development
(a)               Necessary to deal with large amount of yolk in ancestral vertebrate eggs

V.     Animal symmetry

A.    When we examine animals, we note two general forms of “life styles” – either they actively move in specific directions or they don’t

1.                  These differences in “life styles” produce differences in body form

B.     Animals that do not move in specific directions tend to be sessile or pelagic

a)                  Sessile = they don’t move (or move very slowly)

b)                  Pelagic – these animals float in the water column

2.                  Their orientation to the environment isn’t of obvious importance

C.    These animals usually have spherical or radial symmetry (Fig. 9-10)

1.                  Spherical symmetry means that the animal can be divided into mirror images regardless of the orientation of the plane dividing the organism

a)                  Found chiefly in unicellular organisms – rare in multicellular animals

2.                  Radial symmetry occurs when mirror images are obtained when the animal is split into two halves along one axis