THE HEART

A few species of animals can survive without a heart and circulatory system. Some
species of worm manage this by being The Origins of the Major Organs 67 so fl at that all of
their cells come in contact with the environment or are just a few layers away. This allows them
to absorb nutrients directly from their surroundings. Sponges also get along without a heart
because they are perforated by a huge number of pores that deliver seawater to within a
millimeter of every cell. But most organisms have had to solve the problem of providing their
cells with food and oxygen and disposing of wastes through the construction of an intricate
system of internal canals and a pump that keeps fl uid moving along through them. The problem
of supply arises very early in animal development once the organism has become thicker than
about 200 microns, about four times the width of a human hair. At that point all of its cells no
longer have equal access to the environment. The solution is that one of the layers created during
gastrulation—the mesoderm—is transformed into major parts of the circulatory, respiratory, and
digestive systems. As usual, the mesoderm does not manage this alone; it relies on molecular
signals from neighboring tissues. It also requires the development of specialized cell types,
which will be covered in the next chapter. Without a heart, blood would not move through this
complex system; it would stagnate and could not be loaded with fresh oxygen and other
nutrients. It has to be pumped through, and larger animals require more pressure to move it.
Evolution produced various solutions, ranging from simple muscle-lined tubes that squeeze
blood through the body to the complex, four-chambered organs of humans and other mammals.
Comparisons between different types of hearts reveal some of the steps in these transformations.
Each type and developmental stage is a solution to a complex engineering problem. The heart
has to function very early in life to get nutrients to cells. Because the embryo grows so quickly, it
undergoes several major transformations. The circulatory system is a massive work in progress.
New routes are continuously built to supply an evergrowing population of cells. And then there
is the problem of birth, which ejects an embryo that has been living in water into the air. The
entire circulatory system is transformed—within minutes—from a closed “circuit” that feeds off
the mother to one able to cope with oxygen and food. The heart’s origins can be traced back to
events between the second and third weeks of the human embryo’s life, as the neural tube takes
shape. At that point the embryo is stretching to become a long, bumpy oval. Seen in cross-
section, the ectoderm is a bit like one bag containing another bag (the mesoderm); this tissue, in
turn, holds the endoderm and the hollow space that will become the gut. The mesoderm thickens
a bit and opens in the middle, like a slowly infl ating air mattress. Two of these mats develop on
the sides of the notochord (see image on page 70). The space in between will soon become a
tube, formed from mesoderm tissue growing inward from right and left. Before that occurs, a bit
of underlying tissue from the endoderm is squeezed up through the space between the mesoderm
and the notochord. To go back to the air mattress analogy, if the two mattresses are lying on a
carpet (the endoderm) and are pushed together, the carpet might wrinkle between them, making a
bulge. In the organism this bit of tissue will be pinched off. The top of the bulge will develop
into the foregut, which will later become the respiratory system, the stomach, and several other
organs. Now the two mats need to grow together. The mesoderm pushes in from the two sides. It
does not completely fi ll the gap; a hollow space is left in the middle, creating the tube in
between. This leaves the organism with three tubes that grow vertically through the body. Seen
from the top, this has produced the neural tube on the dorsal side, which will develop into the
spinal cord; the foregut in the middle; and the heart on the ventral side. The movements of
endoderm have given the heart tube an inner lining (the endocardium) and an outer one (the
myocardium). As its cells differentiate into muscle, they begin to pulse. Single cells have the
unusual ability to contract by themselves, and they will continue pulsing rhythmically even if
removed from an animal embryo and placed in a test tube. In the animal they fi nd a common
rhythm by communicating with each other by releasing and absorbing two substances, calcium
and The Origins of the Major Organs 69 sodium, in waves. These are the embryo’s fi rst
heartbeats, and they take place long before the heart has developed any familiar structures. It
does not yet pump blood, however; that will start about a week later. The heart of an adult human
is a sac of thick muscle with four inner chambers separated by valves. It is connected to the
circulatory system by large arteries and veins. Building it from a simple tube involves bending
and separating it with valves and septal walls it as it grows. At fi rst, it bulges in some places and
is tightened in others, resembling the relatively simple hearts of fi sh. Later, two major
compartments form, like amphibian hearts. Snakes and turtles have three-chambered hearts. In
some types of aphids the organ is much more complex, with eight. (At least two cases have been
documented of humans with fi vechambered hearts, probably arising through mutations.) At the
beginning of heart formation the embryo has already developed differences between the anterior
and posterior ends and the dorsal and ventral sides. In most people the heart sits slightly to the
left of center. It begins in the middle but by the eighth week has moved to the side. The spleen,
stomach, gut, lungs, and other organs also develop in an asymmetrical way. Their positions are
determined by the behavior of cells in the node very early in development and the activity of
specifi c genes. Imagine placing a long, skinny balloon in a small box and blowing it up: It
would soon run out of room and bend. The same thing happens with the heart; the cardiac tube
quickly outgrows its space and twists into a loop. Cells in different areas begin turning out
specialized proteins. In mice a molecule called Hand1 appears in the left ventricle, while Hand2
is produced only on the right side. In 1998 Deepak Srivastava, a geneticist now at the University
of California at San Francisco, showed that if Hand2 is deleted from mice, the right ventricle is
lost. The left ventricle remains, but it produces only Hand1. When this happens, the heart tube
stops growing. The reason seems to be that Hand proteins interpret asymmetric positional
information in the developing heart and participate in development of segments of the heart tube,
which give rise to specifi c chambers.