Lectures 6&7 Early Development

Embryos are similar across the vertebrates.

Ontogeny Recapitulates Phylogeny Haeckel (1866)

What is Early Development?

For our purposes, early development will be the formation of an embryo from a zygote.

Egg Zygote Sperm

The embryo has 3 germ layers: Ectoderm, Mesoderm, & Endoderm

Embryo

Adult Organism

Plus gut, notochord, coelom, neural tube.

A larva is a free-living embryo able to secure its own nourishment, for example, a tadpole. The early stages of development (to gastrula) occur in all
animals, and are strong evidence for the unity of all animal phyla.

Vertebrate Reproductive Strategies

Vertebrates reproduce sexually (with rare exceptions) and therefore each vertebrate animal must develop from a fertilized egg or zygote. Vertebrate ancestors, and probably early vertebrates as well, would have shed eggs and sperm into the water for fertilization and development to occur in the plankton. Many of these would fuel their own development by feeding on other plankton.

Most living vertebrates retain their eggs or provide them with yolk to allow the embryo to develop to a later stage before having to fend for itself.
There is a tradeoff; more offspring with little yolk and less chance of survival, or fewer offspring with more yolk (or other kinds of parental investment) but better survival.

Egg Types

H&G 5.6, KK 5.2

Egg type according to amount of yolk reflects reproductive strategy. It determines how far the embryo can develop before it needs to feed itself. It also modifies the stages of early development.

Tiny, microlecithal eggs are found in many marine animals with planktonic larvae, e.g. Cephalochordata and Echinodermata. Eggs of Eutheria are secondarily microlecithal. Mesolecithal eggs are found in cyclostomes, some Osteichthyes, Amphibia, and Metatheria (marsupials).
Macrolecithal eggs are found in most fish, Reptilia, Aves and monotremes.

Sagittal, Frontal and Transverse Sections

sagittal section; cuts into left and right halves

transverse section; separates anterior and posterior

frontal section; separates ventral from dorsal

See KK 1.14

transverse frontal

sagittal

Zygote to Blastula in the Microlecithal egg of H&G 5.6, KK 5.2 Amphioxus

site where gastrulation will begin

In microlecithal eggs, early cleavages are radial, complete and equal, leading to a hollow ball of cells called a blastula. The hollow space is called the blastocoel.

Gastrulation and Neurulation in the Microlecithal egg of Amphioxus

sagittal H&G 5.7, KK 5.7, 5.8 transverse

The gastrocoel or embryonic gut develops as an invagination, creating a 2layered embryo or gastrula with ectoderm and endoderm. The third germ layer, or mesoderm, begins on the dorsal part of the endoderm, where the notochord will develop. The cord induces the formation of the neural tube. The embryo is now a neurula.

Amphioxus Neurula

transverse section
Fig. 5.7 H&G

The neural tube is a special kind of ectoderm called neurectodem. An Amphioxus would be a free-swimming (with cilia) and feeding larval stage at this point. The somites would be segmented.

Zygote to Gastrula in the Mesolecithal Egg of an Amphibian KK 5.2, H&G 5.6

In the yolkier egg of an amphibian, the divisions are still complete but markedly unequal. Larger cells are to one side (the vegetal pole), the blastula is several cells thick, and the blastocoel is towards one end (the animal pole).

Amphibian Gastrulation and Neurulation

The infolding process that creates the gastrocoel in the microlecithal egg is highly modified to accommodate all the yolk. Mesoderm formation, again, starts dorsally at the notochord but its internal space occurs by splitting rather than evagination. Again, the notochord induces the neural plate and eventually the neural tube. The mesoderm is partitioned into 3 sections on each side. sagittal transverse

KK 5.11, H&G 5.7

External View

Blastula Formation

Courtesy of Prof. M. Marsden

Gastrulation and Neurulation

Courtesy Prof. M. Marsden

Preview of the fate of the embryonic tissues: 1. ectoderm

Neural crests (unique to vertebrates) will form pharyngeal cartilage and bone, teeth, sensory nerves, acoustic and lateral line system. Neural tube will form the central nervous system.

Ectoderm will produce epidermis and derivatives.

Preview of the fate of the embryonic tissues: 2. mesoderm

Mesenchyme

The outer layer of the epimere, the dermatome, will produce the dermis. The inner layer of the epimere, the myotome, will form the muscles of the trunk. The innermost part will become the schlerotome and move to surround the notochord and become the vertebrae. The mesomere will produce the excretory and reproductive systems.

The hypomere will make the serous lining of the peritoneal cavity, the mesenteries, and the muscles of the gut.

Preview of the fate of the embryonic tissues: 3. endoderm

The endoderm will form the inner layer of the gut, organs of the gut (including lungs) and germ cells.

First Cleavages in a Macrolecithal Egg

The first cell divisions in the macrolecithal egg are partial, and only furrow the surface. These divisions eventually leas to a patch of flat cells called a germinal disk.

The blastula (or blastoderm) is a layer of cells on a large mass of yolk.

Comparison of Blastulas
Blastulas vary in appearance according to the amount of yolk.

Comparison of Gastrulation

Gastrulation in the macrolecithal egg of a bird or reptile looks very different, more of a separation of the germinal disk into two layers than a process of involution. Kardong refers to these layers as epiblast and hypoblast.

Mesoderm Formation in the Bird

KK 5.13, H&G 5.10

Looking down on the flat blastula, it is elongate, and a groove or primitive streak forms at one end.

Cells penetrate at the primitive streak and move out in all directions, creating a middle or mesoderm layer between epiblast (ectoderm) and endoderm.

The origin of the notochord and neural plate

The mesoderm being created directly ahead of the primitive streak will become the notochord. As in other cases, it induces the neural fold above.

KK 5.13, 5.14

Neurulation as seen from above

H&G 5.10 The neural tube forms ahead of the primitive streak, and above the notochord.

With the formation of the neural tube, the embryo takes on a threedimensional shape.

Fate of Extra-embryonic structures

KK 5.13, H&G 5.11 The extra-embryonic mesoderm (amnion) splits to create an extra-embryonic coelomic space. This is called the extra-embryonic coelom. The yolk is enclosed by extraembryonic membrane and a new extraembryonic structure, the allantois, evaginates from the embryos gut. Note that extraembryonic mesoderm is called amnion and extra-embryonic ectoderm is called chorion.

Fate of Extra-embryonic structures 2

.

The allantiois serves as a respiratory organ and as a reservoir for waste in the embryo. The space in which the embryo resides is called the amniotic cavity. As the embryo gets larger, and the yolk smaller, the connection to the yolk becomes a yolk stalk.

Blastula formation in the secondarily microlecithal egg of a eutherian mammal

KK 5.2, H&G 5.6

Eggs of eutherian (placental) mammals are microlecithal. However, their development looks more like other amniotes. There is an inner cell mass of cells that will produce the embryo, while the outer cell layer (trophoblast) will be involved in implantation.

Eutherian Gastrulation
KK Fig. 5.15

As in the bird, the endoderm cells migrate from the ectoderm, and mesoderm arises at a primitive streak (not visible in this transverse section).

Eutherian Neurulation
KK Fig. 5.15

The formation of the primitive streak and mesoderm, including the notochord, is very much like that of a bird or reptile.

Eutherian Extraembryonic Membranes

KK Fig. 5.15

Note: this embryo is rotated 180 degrees compared to the chicken a few slides back.

Although derived from a microlecithal egg, and in a uterus rather than an egg, a eutherian embryo looks rather like other amniote embryos. It is in an amniotic cavity. The yolk sac is vestigial, and it is the chorion that interacts with the uterine wall to make a the placenta. Hence it is a chorioallantoic placenta.