Table of Contents
Fundamentals of Biochemistry
Biochem 380 - Fall 2006
Lecture 003
Outline
- Announcements
- Questions
- Section 5: Biochemistry and Evolution
- Evolution Software Demonstration
- Section 6: The Cell is the Unit of Life
- Section 7: Prokaryotic Cells
- Section 8: Eukaryotic Cells
- Section 9: The Living Cell
Announcements
- Lecture 2 notes are online
- Quiz on Friday on Chapter 1
Questions
- Any questions on the material from the previous lecture?
Section 5: Biochemistry and Evolution
- The theory of evolution, as put forth by Charles Darwin in the 19th century, and subsequently developed
by others, is fundamental for enabling a deep understanding of biochemistry
- The evidence for evolution is overwhelming, and continues to increase over time, as detailed comparisons
of genomic sequences from diverse species reveals common evolutionary origins and histories
- This recent molecular data builds upon and confirms earlier macroscopic support for evolution from
comparative anatomy, population genetics and paleontology
The Origins of Life
- Fossil records indicate the emergence of primitive cellular organisms at least 3 billion years ago
- Commonalities in 'core' metabolic pathways and the near universality of the genetic code indicate that
all modern life descended and diverged from a common ancestor billions of years ago
- Genetic analysis reveals three main branches of modern life: archaebacteria,
eubacteria and eukaryotes
- Both archaebacteria and eubacteria are prokaryotes (lacking a nucleus and other complex cellular features)
while multicellular organisms such as humans have eukaroyotic cells
OOL Theories
- While evolutionary theory is well established, many questions about the details remain to be answered
- Such questions include the emergence of the first replicating systems from pre-living conditions
- Origin-of-Life (OOL) theorists have proposed various theories to address this, such
as Stuart Kauffman's autocatalytic networks, in which a self-sustaining set of chemical reactions
can emerge, with each reaction in the set being catalyzed by the products of one or more of the other reactions
Mechanisms of Evolution
- Once replication is possible, multiple cycles of replication in a population of organisms,
in conjunction with variation and differential rates of survival (selection), can result in large amounts of
change in the phenotypes of the organisms over time
- These processes of reproduction, variation and selection are the fundamental
mechanisms of evolutionary change
- The evolutionary changes in real living organisms is of course vastly more complicated. Recent discoveries
in developmental biology are beginning to shed light on how the variational mechanisms of morphogenesis and cellular
differentiation can contribute to evolutionary change (Evo-Devo book, by Sean Carroll)
Evolution Demo
- The Evolution Demo is a small program that I created to investigate and demonstrate the mechanisms of evolution
- The challenge in creating such a program is two-fold:
- Define a fitness criterion that can be satisfied in a 'non-trivial' manner
(an trivial demonstration of evolution might be the 'monkey typing Shakespeare' scenario)
- At the same time, simulation of the phenotypes must be computationally efficient enough
to demonstrate evolutionary change in real-time
Evolution Demo Implementation
- In this demo, the fitness criterion is to maximize the number of cells in a growing plant
- There are 3 types of cells, and a set of 'growing rules' that are encoded in the plant genome
- With the given rules for simulating a growing plant, a 'good' solution is not immediately obvious
(at least to me)
Evolution Demo
The demo is available for download from the course web pages
Currently, there is only a Windows version
However, the source code is available (any volunteers to port the code to other platforms?)
Survey: who has a computer at home?
What OS do you run? Windows/Mac/Gnu-Linux/Other?
(Run demo)
The Cell is the Unit of Life
- The cell is the smallest system of biomolecules that is considered to be 'alive'
- A cell can be thought of as a droplet of water surrounded by a plasma membrane.
As we will see later, it is actually a droplet of water jam-packed with 'stuff'
- Everything inside the outer membrane, except for the nucleus, is referred to as the
cytoplasm
- The part of the cytoplasm that does not include internal membrane-bound organelles is
called the cytosol
Viruses
- What about viruses? Are they alive? Why or why not?
- Virus are subcellular infectious particles consisting of nucleic acid molecules surrounded by
a protein coat. Although they can reproduce, they require the machinery of a living cell to do so, and therefore
are not considered to be independent living organisms
- The simplicity of viruses makes them useful for understanding biochemistry, however. Viruses
that infect bacteria, called phages (bacteriophages), were important tools for understanding the
genetic machinery of the cell
- Recently, there has been renewed interest in bacteriophages as a possible solution to the growing
problem of antibiotic resistance
Prokaryotic Cells
- Prokaryotes are the simplest kinds of single-celled organisms. Some species have as few as 1000 genes
- Although much simpler than eukaryotes, prokaryotic species are incredibly diverse.
They can be thought of as 'biochemical specialists' that are able to inhabit almost every kind of environment
on Earth
- The most well studied bacterial species is Escherichia coli (E. coli). The biochemistry of this
model organism is used as an example in many parts of the text
Structure of Prokaryotic Cells
- Most bacteria have a rigid cell wall and lack interior membrane compartments
- Many bacteria, including E. coli, also have a second, 'outer' membrane surrounding the cell wall.
This second layer acts as a 'filter', protecting the bacteria from the external environment, while still allowing
nutrients to enter
Eukaryotic Cells
- Eukaryotic organisms can be classified into 4 main groups: plants, animals, fungi and protists
- Eukaryotic cells are more complex than prokaryotic cells. The interior of
a eukaryotic cell has many different kinds of organelles, surrounded and organized by a cytoskeleton
architecture
- The most distinctive organelle is the nucleus, which is tightly packed with the cell's DNA. The nucleus
is also where RNA synthesis and ribosome assembly take place
- Other organelles include the endoplasmic reticulum, the golgi apparatus, mitochondria,
lysosomes and peroxisomes
An Animal Eukaryotic Cell
The Endoplasmic Reticulum
- The endoplasmic reticulum (ER) is a network of membrane sheets and tubes that extend out from the periphery of
the nucleus. The interior of the ER is called the lumen
- The ER serves as a sorting and packaging system for newly-synthesized proteins. Ribosomes attached to
the outside of the ER translate RNA transcripts into new proteins by extruding the proteins across the ER
membrane into the lumen
The Golgi Apparatus
- The Golgi apparatus extend the processing and transport capabilities of the ER through a collection of
sacs and vesicles
-
These are lipid membrane compartments that contain collections of biomolecules for sequential processing, transport,
and in some cases, secretion outside of the cell
The Mitochondrion
- The mitochondrion is another important organelle found in most eukaryotic cells. It is responsible for
aerobic energy production.
It consists of a permeable outer membrane, a highly-folded and largely impermeable inner membrane,
and an interior called the matrix
- Organic compounds are delivered to the mitochondrion, where they are oxidized, releasing electrons.
The electons flow through the inner membrane to combine with O2, transporting protons into the matrix.
These in turn flow back out through ATP Synthetase enzymes, generating ATP
The Living Cell
- Most drawings and pictures fail to capture the incredibly dynamic and beautiful activity of living cells.
A typical cell is packed full of the organelles and macromolecules that we have briefly seen. These structures,
while very small in macroscopic terms, are still huge compared to the scale of atoms and molecules
- Millions of atoms are present within a single macromolecule such as the ribosome. In addition, the
molecules are in constant motion, colliding and diffusing throughout the cell. On average, a typical enzyme
and small molecule will collide with each other around 1 million times per second
- The organization and complexity of molecular traffic within the cytosol is only just beginning to be fully
appreciated. This organization gives the cell a greater degree control of chemical activity, beyond what is possible
through simple solution chemistry
Questions
References
- Book: The Origins of Order, by Stuart Kauffman
A deep and fascinating investigation of evolutionary mechanisms, including
a proposal for 'pre-genomic' autocatalytic metabolic networks
- Book: The Blind Watchmaker, by Richard Dawkins
A popular and very readable exposition of evolutionary theory
- Book: Evo-Devo, by Sean Carroll
A consideration of evolution in the context of developmental biology, including a presentation
of recently-understood mechanisms of morphogenesis and cellular differentiation, and their
potential for contributing to evolutionary change
Next Lecture: Sections 2.1 - 2.3