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As signal compounds, they can also help communicate with other organisms by attracting insects for pollination or animals for seed propagation. Polar secondary compounds are frequently stored in vacuoles, whereas lipophilic compounds are kept in oil vessels, resin channels, or glandular cells. Often, secondary compounds are stored as prodrugs, and only when the plant is wounded or an infection occurs will they be activated – mostly by β‐glucosidase cleaving off a glucose residue. Seeds contain vacuoles storing a protein reserve. Vacuoles can have various functions in a plant cell – providing additional storage space or acting as a defense or signaling compartment. The storage function in particular requires high osmotic pressure inside the vacuole, which is crucial for the stabilization and growth of the plant (turgor regulation). Active transport in plants mostly functions via proton gradients, as opposed to animal cells, which rely more on Na+/K+ gradients, using Na+, K+‐ATPase (see Section 3.1.1.2).
Peroxisomes are small membrane‐enclosed, mostly rounded vesicles in which H2O2 is produced or degraded (e.g. by the enzyme catalase).
3.1.3 Mitochondria and Chloroplasts
Mitochondria are very striking organelles that are found in nearly all eukaryotic cells (Figure 3.14). They look like worms or sausages and are between 1 μm and several micrometers long and 0.5 μm thick. Mitochondria have two separate membrane systems. The inner membrane forms a series of infoldings (cristae) that extend the surface considerably. The large surface area is needed because proteins and enzymes of the respiratory chain need to find space in or on the inner mitochondrial membrane (Figure 3.15). In a liver cell, mitochondria occupy about 22% of the cell volume.
Figure 3.14 Composition of a mitochondrion. (a) Electron microscope photograph.
Source: Courtesy of K.R. Porter/Photo Researchers, Inc.
(b) Schematic representation.
Source: Voet et al. (2016). Adapted with permission of John Wiley and Sons.
Figure 3.15 Function of mitochondrion: metabolism and respiratory chain. (a) Metabolism and respiration in mitochondrion. (b) Schematic representation of the respiratory chain with the complexes I–IV; the proton gradient is used by ATP synthase to produce ATP. Rotenone, malonate, antimycin A, and KCN are the inhibitors of complexes I–IV. FeS, iron–sulfur cluster; Cyt, cytochrome; CoQ, ubiquinone; FMN, flavin mononucleotide.
The respiratory chain produces ATP from the reduction equivalents NADH and FADH2. During this process, electrons are transported through several intermediate stages, and a proton gradient is built up to provide energy for ATP synthase. This is also referred to as cellular respiration because the respiratory chain uses up oxygen. Without mitochondria, aerobic organisms such as animals, fungi, and plants would not be able to use oxygen from the air for the oxidation of organic matter – in other words, to produce energy. There are, however, some bacteria and a few eukaryotes that are anaerobic (i.e. they do not need oxygen). These organisms do not contain mitochondria.
During the citric acid cycle(Krebs cycle), which takes place in the mitochondria, acetyl CoA is introduced, and in each run of the cycle, CO2 and reduction equivalents are generated. The acetyl CoA is derived from pyruvate, a product of glycolysis, which has been taken up by the mitochondria through a pyruvate transporter. It is then converted into acetyl CoA by a pyruvate decarboxylase complex. Another way of generating acetyl CoA is by β‐oxidation of fatty acids – a process that also takes place in mitochondria (Figure 3.15).
Mitochondria contain their own ring‐shaped DNA (Figure 3.16). In animals, the mitochondrial genome (mtDNA) is significantly smaller (16–19 kb) than in plants. It contains 13 genes coding for enzymes or other proteins involved in electron transport, 22 genes for tRNAs, and two genes for rRNAs. As every animal cell contains several hundred or even thousand mitochondria, each of which contains 5–10 mtDNA copies, the total of mtDNA copies amounts to several thousand per cell. mtDNA makes up about 1% of the total amount of DNA contained in a cell. The analysis of nucleotide sequences from mitochondrial genes has become an important tool in systematics to establish phylogenies and to define species.
Figure 3.16 Schematic overview of the arrangement of genes in the mtDNA of mammals.
Plant mitochondria, by contrast, have large genomes (150–2500 kb). Some of their genes even have an intron/exon structure.
Mitochondria contain functional ribosomes equivalent to the prokaryotic 70S type, and the nucleotide sequences in mitochondrial genes and the amino acid sequences of the respective proteins are more closely related to the corresponding prokaryotic genes than to equivalents coded in the nucleus. The genetic code of mitochondria shows a few differences to the universal code: UGA (stop codon) codes in animals and fungi for tryptophan, AUA (for isoleucine) codes in animals and fungi for methionine, and AGG (arginine) codes in mammals for stop and in invertebrates for serine.
These findings as well as other mitochondrial characteristics led to the endosymbiont hypothesis, which states that mitochondria are derived from α‐purple bacteria that were ingested by an ancestral eucyte 1.2 billion years ago and lived on as endosymbionts. The cell provides nutrients for the endosymbionts and receives ATP in return. Figure 3.17 shows a likely ingestion path for the α‐purple bacteria into the ancestral eucyte. It is assumed that the ancestral eucyte came into being by the infolding of a bacterial cytoplasmic membrane to form an ER. The membrane then began to surround the chromosome, thus forming a nucleus.
Figure 3.17 Development of an early eucyte and origin of mitochondria. α‐Purple bacteria were ingested by the early eucyte in a kind of phagocytosis. Hence, the outer mitochondrial membrane is derived from the host cell, whereas the inner mitochondrial membrane is the original bacterial cytoplasmic membrane.
Green plants and algae contain an additional organelle, the conspicuous chloroplasts, which are significantly larger and structurally more complex than mitochondria (