Germ layer

A germ layer is a primary layer of cells that form during embryogenesis.[1] The three
germ layers in vertebrates are particularly pronounced; however, all eumetazoans,
(animalsmore complex than the sponge) produce two or three primary germ layers.
Animals with radial symmetry, like cnidarians, produce two germ layers
(the ectoderm and endoderm) making them diploblastic. Animals with bilateral
symmetry produce a third layer (the mesoderm), between these two layers. making
them triploblastic. Germ layers eventually give rise to all of an
animal’s tissues and organs through the process of organogenesis.

A germ layer is a group of cells in an embryo that interact with each other as the embryo
develops and contribute to the formation of all organs and tissues. All animals, except
perhaps sponges, form two or three germ layers. The germ layers develop early in
embryonic life, through the process of gastrulation. During gastrulation, a hollow cluster
of cells called a blastula reorganizes into two primary germ layers: an inner layer,
called endoderm, and an outer layer, called ectoderm. Diploblastic organisms have only
the two primary germ layers; these organisms characteristically have multiple
symmetrical body axes (radial symmetry), as is true of jellyfish, sea anemones, and the
rest of the phylum Cnidaria. All other animals are triploblastic,
as endoderm and ectoderm interact to produce a third germ layer, called mesoderm.
Together, the three germ layers will give rise to every organ in the body, from skin and
hair to the digestive tract.

Gastrulation differs across species, but the general process is the same: the hollow ball
of cells that forms the blastula differentiates into layers. The first phase
of gastrulation produces a two-layered organism comprised of ectoderm and endoderm.
The ectoderm will form the outer components of the body, such as skin, hair, and
mammary glands, as well as part of the nervous system. Following gastrulation, a
section of the ectoderm folds inward, creating a groove that closes and forms an
isolated tube down the dorsal midsection of the embryo. This process
of neurulation forms the neural tube, which gives rise to the central nervous system.
During neurulation, ectoderm also forms a type of tissue called the neural crest, which
helps to form structures of the face and brain. The endoderm produced
during gastrulation will form the lining of the digestive tract, as well as that of the lungs
and thyroid. For animals with three germ layers, after the endoderm and ectoderm have
formed, interactions between the two germ layers induce the development
of mesoderm. The mesoderm forms skeletal muscle, bone, connective tissue, the heart,
and the urogenital system. Due to the evolution of the mesoderm, triploblastic animals
develop visceral organs such as stomachs and intestines, rather than retaining the open
digestive cavity characteristic of diploblastic animals.

Christian Pander, a doctoral student of Ignaz Döllinger at the University of Würzburg, in

Würzburg, Germany, first recognized the existence of germ layers in chicks (Gallus
gallus) in 1817. In the publications derived from his dissertation, Pander described how
two layers of cells, which he called serous and mucous, gave rise to an intermediate
layer, which he called vascular. Pander wrote of the interdependence of these three
layers as well as the necessity of their interaction to form organs.

In 1825, eight years after Pander's initial descriptions, Martin Rathke, a physician and
embryologist from Prussia (now Poland), discovered layers of cells in a developing
invertebrate crayfish, Astacus astacus, that corresponded to those Pander had
described in chicks. Rathke showed that the embryonic layers Pander described existed
in animals outside of the vertebrate clade. Karl Ernst von Baer, a professor of anatomy
at the University of Königsberg, in Königsberg, Germany, applied Pander's germ layer
concept to all vertebrates in his 1828 Über Entwicklungsgeschichte der Thiere.
Beobachtung und Reflexion (On the Developmental History of the Animals.
Observations and Reflections).

Discussion of the germ layers dwindled over the next twenty one years, but they
resurfaced when Thomas Henry Huxley, a natural historian in England, published "On
the Anatomy and Affinities of the Family of the Medusae." In that 1849 text, Huxley
suggested that adult jellyfish (Medusae) possessed two tissue layers, which he called
foundation membranes, that relate to each other in the same manner that Pander had
observed of the serous and mucous layers in the chick embryo. Huxley realized that a
correlation existed between the body architecture of the adult jellyfish and the vertebrate
embryo. Based on that association, Huxley attempted to integrate the study of
vertebrates with that of invertebrates, and to unite studies of development, or ontogeny,
with studies of organismal relationships, or phylogeny. The relationship
between ontogeny and phylogeny, later called recapitulation, would be adopted and
expanded by proponents of evolution, including Charles Darwin, in England, and Ernst
Haeckel, a professor of comparative anatomy at the University of Jena, in Jena,
Germany.

In the six years following Huxley's publication on Medusae, embryologist Robert Remak,
in Germany, refined germ layer theory in two ways in his treatise Untersuchungen über
die Bildung und Entwickelung der Wirbelthiere (Studies on the Formation and
Development of Vertebrates). First, while working as a microscopist, Remak noticed
that all of the germ layer cells of the chick embryo derived from the original single cell of
the fertilized egg. Thus, Remak concluded, all cells originate from division of pre-
existing cells, a conclusion that became central to cell theory. Second, Remak provided
histological evidence for the existence of three distinct germ layers and traced the
derivatives of each throughout chick development. Few noticed Remak's contributions
to cell theory and research on germ layers.

In 1867 Aleksandr Kovalevsky, professor of embryology at the University of St.

Petersburg, in St. Petersburg, Russia, published a series of studies that showed the
presence of germ layers among invertebrates. Kovalevsky's work established the
universality and homologous nature of the germ layers within the animal kingdom.

According to Jane Oppenheimer, a biologist and historian of science who worked

at Bryn Mawr College in Philadelphia, Pennsylvania during the twentieth century,
Kovalevsky's research prompted some of the most prominent scientists of the
nineteenth century to research on the germ layers. The concept of the germ layers as
invariant across species soon became entrenched and formed the basis of germ layer
theory. Germ layer theory held that each of the germ layers, regardless of species, gave
rise to a fixed set of organs. In 1872 Ernst Haeckelcombined observations of germ
layers with evolutionary theory to hypothesize that an unknown two-layered organism,
which he called a gastraea, was ancestral to all other animals; this came to be known
as the Gastraea Theory. One year later, Edwin Ray Lankester, Professor of Zoology
at University College, London, in London, England, published a theory similar to
Haeckel's along with a classification of all animals based on their composition of germ
layers: homoblastic, diploblastic, and triploblastic. Researchers still use Lankester's
classification.

In the late 1870s, several years after Haeckel's and Lankester's publications, many
embryologists challenged germ layer theory and Haeckel's Gastraea theory. Wilhelm
His, Rudolf Albert von Kölliker, and Oscar and Richard Hertwig, all in Germany at the
time, objected to the germ layer theory. In a series of publications from 1878 through
1881, the Hertwig brothers provided evidence that the germ layers had greater
capacities for differentiation than most scientists recognized. In 1881 the Hertwigs
formulated their Coelom theory, which focused on the role of mesoderm and also
introduced the term and concept of mesenchyme, a type of animal tissue derived mostly
from mesoderm.

Amid the varied arguments supporting or denying germ layer theory, some
embryologists in the 1890s began to refocus their efforts on methods that could help
them further understand how animals develop, and they employed physical
manipulations of embryos rather than purely observational or descriptive embryology. In
1901 Charles Sedgwick Minot, a professor at Harvard Medical School in Boston,
Massachusetts, predicted that the transplantation of cells from one germ layer onto
another resulted in those cells adopting the fate of their new environment.

More than twenty years later, in 1924, Hilde Proescholdt Mangold and her doctoral
advisor at the Zoological Institute in Freiburg, Germany, Hans Spemann, found
evidence for Minot's prediction and dismantled the foundation of the germ layer theory.
Mangold harvested presumptive ectoderm from the dorsal lip, a tissue that organizes
the gastrula stage, of an embryonic newt and transplanted this tissue to a different germ
layer of the gastrula of a second species of newt. The transplanted ectoderm responded
to the local environment on the developing host newt, and induced the formation of an
extra head, nervous system structure, or extra body. That experiment demonstrated that
the fates of germ layer cells are not entirely predetermined at the start of development.

In the fifteen years following Mangold's work, embryologists continued to explore the
potential for the three germ layers to differentiate along different routes and they
produced evidence that further weakened the germ layer theory. Sven Hörstadius,
professor at Uppsala University, in Uppsala, Sweden, used echinoderms, such as sea
urchins, to study how germ layers differentiate. He employed transplantation,
recombination, and fate mapping experiments to investigate the capacity of the germ
layers to transform into tissues atypical of normal differentiation.

Throughout the remainder of the twentieth century, researchers continued to

accumulate evidence that invalidated the theory that germ layers are pre-defined or
highly-fated tissues. Following the works of Spemann, Mangold, and Hörstadius,
scientists further explored germ layer potential for varied development. In the early
1950s Robert Briggs, at Indiana University in Bloomington, Indiana, and Thomas King,
at the Institute for Cancer Research in Philadelphia, Pennsylvania, transplanted nuclei
from the presumptive endoderm of the northern leopard frog, Rana pipiens, into eggs
from which they had removed the nuclei. Briggs and King tracked the development of
these transplanted nuclei to explore the timing of cell differentiation, and with those
experiments they laid the foundation for future research in cloning. In the late 1960s
Pieter D. Nieuwkoop, at the Hubrecht Laboratory in the Royal Netherlands Academy of
Arts and Science, in Utrecht, Holland, discovered that endoderm induces
adjacent ectoderm to form mesoderm. In the 1980s scientists shifted their focus towards
identifying the genes that induce structural differentiation of the germ layers.
Researchers in the early twenty-first century investigated the regulatory networks
through which individual genes interact to cause germ layer differentiation.
Germ layers

Gastrulation of a diploblast: The formation of germ layers from a (1) blastula to a (2)
gastrula. Some of the ectoderm cells (orange) move inward forming the endoderm (red).

Micrograph of a teratoma, a tumour that characteristically has tissue from all three germ
layers. The image shows tissue derived from the mesoderm (immature cartilage - left-
upper corner of image), endoderm(gastrointestinal glands - center-bottom of image)
and ectoderm (epidermis - right of image). H&E stain.
Caspar Friedrich Wolff observed organization of the early embryo in leaf-like layers. In
1817, Heinz Christian Panderdiscovered three primordial germ layers while studying
chick embryos. Between 1850 and 1855, Robert Remak had further refined the germ
cell layer concept, and introduced into English were the terms "mesoderm" by Huxley in
1871 and "ectoderm" and "endoderm" by Lankester in 1873.
Among animals, sponges show the simplest organization, having a single germ layer.
Although they have differentiated cells(e.g. collar cells), they lack true tissue
coordination. Diploblastic animals, Cnidaria and Ctenophora, show an increase in
complexity, having two germ layers, the endoderm and ectoderm. Diploblastic animals
are organized into recognisable tissues. All higher animals (from flatworms to humans)
are triploblastic, possessing a mesoderm in addition to the germ layers found in
Diploblasts. Triploblastic animals develop recognizable organs.

Development
Fertilization leads to the formation of a zygote. During the next
stage, cleavage, mitotic cell divisions transform the zygote into a hollow ball of cells,
a blastula. This early embryonic form undergoes gastrulation, forming a gastrula with
either two or three layers (the germ layers). In all vertebrates, these progenitor cells
differentiate into all adult tissues and organs.[2]
In humans, after about three days, the zygote forms a solid mass of cells by mitotic
division, called a morula. This then changes to a blastocyst, consisting of an outer layer
called a trophoblast, and an inner cell mass called the embryoblast. Filled with uterine
fluid, the blastocyst breaks out of the zona pellucida and undergoes implantation. The
inner cell mass initially has two layers: the hypoblast and epiblast. At the end of the
second week, a primitive streak appears. The epiblast in this region moves towards the
primitive streak, dives down into it, and forms a new layer, called the endoderm,
pushing the hypoblast out of the way (this goes on to form the amnion.) The epiblast
keeps moving and forms a second layer, the mesoderm. The top layer is now called
the ectoderm.[3]

Endoderm

The endoderm produces tissue within the lungs, thyroid, and pancreas.
The endoderm is one of the germ layers formed during animal embryogenesis. Cells
migrating inward along the archenteron form the inner layer of the gastrula, which
develops into the endoderm.
The endoderm consists at first of flattened cells, which subsequently become columnar.
It forms the epithelial lining of the whole of the digestive tube except part of the mouth
and pharynx and the terminal part of the rectum (which are lined by involutions of the
ectoderm). It also forms the lining cells of all the glands which open into the digestive
tube, including those of the liver and pancreas; the epithelium of the auditory tube and
tympanic cavity; the trachea, bronchi, and air cells of the lungs; the urinary bladder and
part of the urethra; and the follicle lining of the thyroid gland and thymus.
The endoderm forms: the stomach, the colon, the liver, the pancreas, the urinary
bladder, the epithelial parts of trachea, the lungs, the pharynx, the thyroid, the
parathyroid, and the intestines.

Mesoderm

The mesoderm aids in the production of cardiac muscle, skeletal muscle, smooth
muscle, tissues within the kidneys, and red blood cells.
The mesoderm germ layer forms in the embryos of triploblastic animals.
During gastrulation, some of the cells migrating inward contribute to the mesoderm, an
additional layer between the endoderm and the ectoderm.[citation needed] The formation of a
mesoderm leads to the development of a coelom. Organs formed inside a coelom can
freely move, grow, and develop independently of the body wall while fluid cushions and
protects them from shocks.[citation needed]
The mesoderm has several components which develop into tissues: intermediate
mesoderm, paraxial mesoderm, lateral plate mesoderm, and chorda-mesoderm. The
chorda-mesoderm develops into the notochord. The intermediate mesoderm develops
into kidneys and gonads. The paraxial mesoderm develops into cartilage, skeletal
muscle, and dermis. The lateral plate mesodermdevelops into the circulatory system
(including the heart and spleen), the wall of the gut, and wall of the human body. [4]
Through cell signaling cascades and interactions with the ectodermal and endodermal
cells, the mesodermal cells begin the process of differentiation.[5]
The mesoderm forms: muscle (smooth and striated), bone, cartilage, connective
tissue, adipose tissue, circulatory system, lymphatic system, dermis, genitourinary
system, serous membranes, and notochord.

Ectoderm

The ectoderm produces tissues within the epidermis, aids in the formation
of neurons within the brain, and constructs melanocytes.
The ectoderm generates the outer layer of the embryo, and it forms from the
embryo's epiblast.[6] The ectoderm develops into the surface ectoderm, neural crest,
and the neural tube.[7]
The surface ectoderm develops into: epidermis, hair, nails, lens of the eye, sebaceous
glands, cornea, tooth enamel, the epithelium of the mouth and nose.
The neural crest of the ectoderm develops into: peripheral nervous system, adrenal
medulla, melanocytes, facial cartilage, dentin of teeth.
The neural tube of the ectoderm develops into: brain, spinal cord, posterior
pituitary, motor neurons, retina.
Note: The anterior pituitary develops from the ectodermal tissue of Rathke's pouch.

Neural crest
Because of its great importance, the neural crest is sometimes considered a fourth
germ layer.[8] It is, however, derived from the ectoderm.

What causes a plant to increase in diameter, or girth? Describe how it

happen and cite the tissues involved?
Vascular Cambium and Cork Cambium are two lateral meristems (undifferentiated cells)
that are responsible for the secondary growth of the plant. Lateral meristems produce tissues
that increase the diameter/girth of the plant.
 OUTPUT
IN
GENERAL
BIOLOGY

Submitted by:
Louie Ann J. Villaraza

Submitted to:
Ms. Ivy Diane Mangampo