Section 1: DNA as the Primary Genetic
Material
Early
Speculations on Genetic Material In the 1920s, it was
already understood that proteins are biological macromolecules composed of
various amino acids linked together. The different sequences of amino acids
could form different proteins, leading people to naturally speculate that the
diverse arrangements of amino acids might contain genetic information. At the
time, studies on other biological macromolecules had not yet revealed similar
structural characteristics. Consequently, most scientists believed that
proteins were the genetic material of living organisms. It wasn't until the 1930s
that it became known that DNA is a biological macromolecule composed of many
deoxyribonucleotides, with the chemical composition of these nucleotides
including phosphate, bases, and deoxyribose (Figure 3-1). There are four types
of deoxyribonucleotides that make up DNA, each with a specific base. This
understanding could have highlighted the importance of DNA, but due to the lack
of clear knowledge about its structure, the view that proteins were the genetic
material still predominated. Griffith's
Transformation Experiment The first challenge to the
idea that proteins were the genetic material, supported by solid experimental
evidence, came from American microbiologist Oswald Avery (O. Avery, 1877–1955),
building on the earlier work of British microbiologist Frederick Griffith (F.
Griffith, 1877–1941). In 1928, Griffith used mice
as experimental subjects to study the pathogenicity of Streptococcus
pneumoniae (pneumococcus). He infected mice with two different types of Streptococcus
pneumoniae. One type had a polysaccharide capsule and formed smooth
colonies on culture media, known as S-type bacteria. S-type bacteria are
pathogenic and can cause pneumonia in humans and mice, leading to septicemia
and death in mice. The other type lacked the polysaccharide capsule and formed
rough colonies on culture media, known as R-type bacteria. R-type bacteria do
not cause disease in humans or mice, and thus are non-pathogenic. Griffith's
experimental process is shown in Figure 3-2. From the bodies of mice in
the fourth group of experiments, live S-type bacteria were isolated, and their
offspring were also pathogenic S-type bacteria. From this, it could be inferred
that the heat-killed S-type bacteria contained some active substance—referred
to as the "transforming factor"—that caused the live R-type bacteria
to transform into live S-type bacteria. But what exactly was this
transforming factor? In the 1940s, Avery and his
colleagues crushed heat-killed S-type bacteria and attempted to remove most of
the sugars, proteins, and lipids, creating a cell extract. When this extract
was added to a culture medium containing live R-type bacteria, live S-type
bacteria appeared (Figure 3-3, Group 1). Then, they treated the cell extract
with different enzymes before conducting the transformation experiment again.
The results showed that the cell extract retained its transforming activity
after treatment with protease, RNase, or esterase (Figure 3-3, Groups 2 to 4),
but lost its transforming activity after treatment with DNase (Figure 3-3,
Group 5). These experiments
demonstrated that the cell extract contained the transforming factor described
earlier, and this transforming factor was likely DNA. Avery and his team
further analyzed the physicochemical properties of the cell extract and found
that they were very similar to those of DNA. As a result, Avery proposed a
conclusion that differed from the prevailing view of most scientists at the
time: DNA is the substance that causes stable genetic changes in R-type
bacteria. The
Hershey-Chase Experiment In 1952, American
geneticist Alfred Hershey (A.D. Hershey, 1908–1997) and his assistant Martha
Chase (M.C. Chase, 1927–2003) conducted another compelling experiment using T2
bacteriophage (Figure 3-4) as their experimental material, utilizing
radioactive isotope labeling technology. The T2 bacteriophage is a
virus that specifically parasitizes within Escherichia coli (E. coli).
Its head and tail are both composed of protein, with DNA inside the head. When
the T2 bacteriophage infects E. coli (Figure 3-5), it uses the
materials within the E. coli cell to synthesize its own components and
undergoes extensive replication under the influence of its own genetic
material. Once the bacteriophages have proliferated to a certain number, the E.
coli cell lyses, releasing a large number of bacteriophages. Hershey and Chase first
cultured E. coli in media containing radioactive isotopes S35 and P32,
respectively, and then used these E. coli to culture T2
bacteriophages. This resulted in bacteriophages with protein labeled with S35
or DNA labeled with P32. They then used the S35 or P32 labeled T2
bacteriophages to infect unlabeled E. coli. After a short period of
incubation, they used a blender to agitate and then centrifuge the mixture
(Figure 3-6). The purpose of agitation was to separate the bacteriophages from
the bacteria, and centrifugation was to precipitate the heavier E. coli
cells, leaving the lighter T2 bacteriophage particles in the supernatant. After
centrifugation, they checked for the presence of radioactive material in the
supernatant and the precipitate. They found that in the infection experiment
with S35-labeled bacteriophages, the radioactive isotopes were primarily found
in the supernatant. In the experiment with P32-labeled bacteriophages, the
radioactive isotopes were primarily found in the precipitate. What does this
result indicate? Further observation
revealed that the bacteriophages released by the lysed bacteria contained
P32-labeled DNA but no S35-labeled protein. What does this result suggest? Hershey and Chase's
experiment demonstrated that when bacteriophages infect bacteria, DNA enters
the bacterial cell while the protein coat remains outside the cell. Therefore,
the characteristics of progeny bacteriophages are inherited through the
parental DNA. DNA is the genetic material of the bacteriophage. Is
DNA the Only Genetic Material? Subsequent research has
shown that genetic material is not limited to DNA; RNA can also serve as
genetic material. Some viruses contain only protein and RNA, such as the
tobacco mosaic virus (Figure 3-7). Protein extracted from the tobacco mosaic
virus cannot cause tobacco plants to become infected, but RNA extracted from
these viruses can. Therefore, in these viruses, RNA is the genetic material.
However, since the vast majority of organisms use DNA as their genetic
material, DNA is considered the primary genetic material. |
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