Is A Chloroplast Found In A Plant Or Animal Cell
Med Sci Monit. 2015; 21: 2073–2078.
Mitochondria, Chloroplasts in Animal and Plant Cells: Significance of Conformational Matching
Received 2015 May 25; Accustomed 2015 Jun 26.
Abstract
Many commonalities betwixt chloroplasts and mitochondria exist, thereby suggesting a common origin via a bacterial ancestor capable of enhanced ATP-dependent free energy product functionally linked to cellular respiration and photosynthesis. Accordingly, the molecular development/retention of the catalytic Qo quinol oxidation site of cytochrome b complexes as the tetrapeptide PEWY sequence functionally underlies the common retention of a chemiosmotic proton gradient mechanism for ATP synthesis in cellular respiration and photosynthesis. Furthermore, the dual regulatory targeting of mitochondrial and chloroplast gene expression by mitochondrial transcription termination factor (MTERF) proteins to promote optimal energy production and oxygen consumption further advances these evolutionary contentions. As a functional effect of enhanced oxygen utilization and production, significant levels of reactive oxygen species (ROS) may exist generated within mitochondria and chloroplasts, which may finer compromise cellular energy product following prolonged stress/inflammationary conditions. Interestingly, both types of organelles accept been identified in selected animal cells, most notably specialized digestive cells lining the gut of several species of Sacoglossan sea slugs. Termed kleptoplasty or kleptoplastic endosymbiosis, functional chloroplasts from algal food sources are internalized and stored within digestive cells to provide the host with dual energy sources derived from mitochondrial and photosynthetic processes. Recently, the ascertainment of internalized algae within embryonic tissues of the spotted salamander strongly suggest that developmental processes within a vertebrate organism may require photosynthetic endosymbiosis every bit an internal regulator. The dual presence of mitochondria and functional chloroplasts within specialized animate being cells indicates a loftier degree of biochemical identity, stereoselectivity, and conformational matching that are the likely keys to their functional presence and essential endosymbiotic activities for over 2.5 billion years.
MeSH Keywords: Chloroplasts, Kleptoplasty, Mitochondria, MTERF, PEWY, Reactive Oxygen Species, Stereospecificity
Background
Mitochondria and chloroplasts represent endosymbiont models of circuitous organelle development driven by evolutionary modification of permanently enslaved primordial bacteria[1–4]. Over diverse eukaryotic phyla mitochondria and chloroplasts either lone or together provide a concerted distension of cellular energy production via shared biochemical pathways. Cellular dysregulation of these 2 distinct organelles may generate potentially dangerous reactive oxygen species (ROS) due to compromised complex bioenergetics energy production, systemic oxidative stress and compounded pro-inflammatory processes. Importantly, genetically- or biochemically-mediated failure of mitochondrial function in human populations represents a potentially dire cistron in the etiology of major disease states that include Type 2 diabetes, atherosclerosis, rheumatoid arthritis, Alzheimer's Illness, and cancer progression [5–21]. In sum, these compelling mechanistic and clinical data suggest that the extent of mitochondrial/chloroplast regulatory signaling may vary over the lifetime of the eukaryotic cell according to physiological demand and bioenergetics requirements[22,23].
Interestingly, a tumor cell may exist viewed as a phenotypic reversion to the last common eukaryotic ancestor of the host cell, i.east., a facultative anaerobic microbe with unlimited replication potential [24]. For example, anaerobic mitochondria in gill cilia of Grand. edulis have evolved to utilize the phenotype of a facultative anaerobe, demonstrating that this primitive type of respiration has been evolutionarily conserved [25,26]. Accordingly, anaerobically functioning mitochondria may represent a re-emergence or evolutionary retrofit of primordial metabolic processes.
It has become recently apparent that mitochondria have discrete microenvironments composed of complex intracellular membrane structures with distinct functional identities determined past segregated biochemical pathways [27] (Figure 1). Given the shared chemical messengers between the 2 and interrelationships between the common energy processes it is not surprising that additional commonalities are emerging. Furthermore, it is no surprise that mitochondria are present in both plants and animals, implying major shared regulatory, bioenergetic, and chemic substrate pathways. Commonalities of free energy processing in both plants and animals have become even stronger by the finding that chloroplast can exist plant in fauna cells. The discovery of kleptoplasty, a functional chloroplast in cells of a non-photosynthetic host [28] is a remarkable miracle [28–31]. It is also institute in metazoans, i.east., the sacoglossan sea slug. Of equal importance is the longevity of functional kleptoplasts in the host, suggesting over again the common significance of bidirectional communication and the many commonalities in molecules exist and then that this phenomenon tin take identify and work. These sea slugs extract and comprise functional chloroplasts from Ulvophyceae into their gut cells [32], allowing their derived "food" to exist gained for months. The dependence on specific algae strongly suggests mutual bidirectional communication is responsible for these phenomena.
The prokaryotic cell is characterized by a general lack of highly structured intracellular organelles but displays intracellular regions of functionality with some membrane enhancements, east.1000., mesosome. We surmise that with time this relatively uncomplicated structure became more than elaborate, adding membrane expanse to perform work, enhancing a major role like respiration. In all probability the stimulus was solar free energy, causing the photolysis of water. This cell was driven in this management because it provided a new coping strategy for, counter-intuitively, DNA advancement. This evolving cellular compages could not survive on its own given the presence of past products it produced, due east.g., ROS, which are basically toxic to unprotected intracellular components, notably Deoxyribonucleic acid. This evolutionary self protection mechanism was further advanced when oxygen levels increased every bit a event of photosynthesis. In all probability the cellular oxygen toxicity issue was partly solved past having a "bacterium" develop in a "bacterium", becoming a eukaryotic jail cell, which could harvest specific bond energy. This also aided in ROS protection with a more structured and protected environment for this new intracellular relationship to evolve, having a plentiful free energy supply for novel DNA expression. Accordingly, a major free radical and gratis radical creator was effectively removed via chloroplasts, which originated in a similar mode as mitochondria. Thus, it is non surprising to find both types of "leaner" in the aforementioned cell and others where only 1 is nowadays. Furthermore, given this shut evolvement, enslavement was not an result in this circumstance because each "cell" used the same or like chemical messengers, stabilizing what appears to be a precarious relationship. Indeed, bidirectional communication served as the process for eukaryotic cellular communication/cooperation, which immune for metazoan evolution. Interestingly, metazoan development is even so highly dependent on the intracellular communication with its endogenous bacterial components from which information technology evolved, e.g., intracellular and extracellular (gut microbiome). The vulnerability expresses itself in "mitochondrial dysfunction" in that information technology tin be so complicated and diverse depending on the tissue region affected. We further surmise that hypoxia plays a major office in triggering mitochondrial dysfunction since this entire relationship depends on a continuously ongoing energy processing system[ii,21]. Briefly, the evolutionary advancement of eukaryotic cells requires this homeostatic energy balance to maintain its multicomponent and faceted existence. Any deviation from the thermodynamically stabilized life form creates a pathology wherever it occurs. This process may besides represent the deleterious mechanisms that may exist associated with aging.
The power of a chloroplast to function as a symbiotic bioenergetic organelle within the intracellular milieu of a representative invertebrate, i.e., the Sacoglossan sea slug, was previously identified as a unique phenomenon unlikely to occur in vertebrates [28–32]. Recently, the observation of internalized algae inside embryonic tissues of the spotted salamander strongly suggest that developmental processes within a vertebrate organism may crave photosynthetic endosymbiosis as an internal regulator [33]. Accordingly, it appears that green algae and spotted salamander embryos have an intimate endosymbiotic relationship and algae are able to invade the embryonic tissues and cells of the salamander and somewhen dethrone as the larvae develop over time [33]. Although endosymbiotic algal cells go through degradation, the cells can as well encyst on the inner capsule wall which is detected through 18s rDNA amplification in the reproductive tracts of the adult salamanders, thereby allowing for the transfer of genes from ane generation to the side by side [33]. Due to the dense accumulation of algae within the embryo, a distinct light-green color is exhibited which leads to beneficial effects for the embryo. Requisite physiological effects include lowering embryonic bloodshed, larger embryo size, and earlier hatching times. Information technology is withal unclear if the algae and the embryo have a truthful bidirectional symbiotic relationship because there is evidence that the algae have no increment in oxygen levels, simply they may do good from the embryos when their nitrogenous waste matter is released. In whatever result, this phenomenon defines a distinctive relationship betwixt developmental processes in a defined vertebrate organism and eukaryotic algae.
A careful examination of the biomedical literature has yielded many examples of existential commonalities between mitochondria and chloroplasts, which include free living bacteria [34]. Formally known as the PEWY motif in mitochondrial complexes, cyt b displays four tetrapeptide residues (PDWY, PPWF, PVWY and PEWY) employed in catalytic reactions, which is now identified every bit the Qo motif. PEWY, which is present in chloroplasts and mitochondria, and PDWY which is present in Gram-positive bacteria both associate with the redox potential of quinone species [34]. These data suggest that when electron transfer occurs from a low-high potential throughout evolution that the cyt bc1 complex with PEWY beingness the Qo motif will function best with a loftier potential and ubiquinone equally its substrate [34]. For PDWY as the cyt b complex, a depression potential and menaquinone volition part the best. In sum, the molecular evolution/retention of the catalytic Qo quinol oxidation site of cytochrome b complexes, functionally underlies the mutual memory of a chemiosmotic proton gradient mechanism for ATP synthesis in cellular respiration and photosynthesis.
The relationship betwixt photosynthesis and respiration tin can vary, thereby demonstrating their dynamic nature. For case, when tomato fruit ripen, their chloroplasts will alter into photosynthetically inactive chromoplasts that tin produce ATP through a respiration procedure known as chromorespiration [35]. Oxygen consumption through chromorespiration tin exist stimulated by NADH and NADPH, and is also sensitive to the plastidial terminal oxidase inhibitor octyl gallate. Isolated chromoplasts are also sensitive to multiple molecules such as the cytochrome b six f complex inhibitor two,5-dibromo-iii-methyl-6-isoproply-p-benzoquinone [35]. Cytochrome f was identified in the chromoplast as was cytochrome c6 and their expression increases in ripened tomatoes suggesting that they may be interim as electron acceptors for the cytochrome b 6 f complex. During ripening, mitochondrial numbers significantly decrease in the fruit tissue [35]. In lodge to recoup for this strong decrease, the number of chromoplasts will functionally increment during the afterward stages of ripening, thereby demonstrating critical modification of energy processing.
Importantly, plants require imported oxygen to acquit out most of their biochemical reactions such as respiration even though they lack the power to distribute oxygen to the cells [36]. To recoup for the lack of this distribution machinery, plants often display steep oxygen gradients that may be impaired due to ecology distress [36]. Thus, plants require different physiological responses to manage the variations of oxygen levels bachelor to them and display metabolic adaptations in energy requirements. Every bit a key example, physiological demand is coupled to activation of the cellular glycolytic pathway to generate ATP production when oxidative phosphorylation is compromised [27]. Cellular oxygen levels have been demonstrated to regulate the expression of Group-VII ethylene response factors (ERFs), a family of transcription factors involved in the regulation of hypoxia-inducible genes that include HRE1 and HRE2 [36]. Furthermore, the functional integrity of mitochondria and chloroplasts are critically linked to cellular oxygen requirements, as regulated by the N-end rule signaling pathway due to the impacted loss. The N-stop rule signaling pathway represents a cellular response mechanism that requires ubiquitin ligation linked to proteasomal degradation via covalent modification of N-terminal amino acids [36].
Finally, the array of complex control machinery by which organellar factor expression (OGE) promotes respiration, photosynthesis and plant development is actively nether investigation [37]. Presently, several required components take been identified that have been functionally associated with OGE processes. Nucleus-encoded proteins have important roles in OGE by promoting various required functions such as splicing, transcription, RNA processing and regulation of translational processes. Normative OGE is regulated by the family unit of mitochondrial transcription termination factors (mTERF). Mammalian mTERFS were originally proposed to specifically end transcription, merely further biochemical and molecular studies indicate that three out of the four mTERFS possess of import regulatory activities necessary for ribosomal biogenesis and antisense transcription termination. Approximately 30 members of the mTERF family unit have been identified throughout constitute evolution, merely withal little is known about how photosynthetic organisms are using mTERFs and OGE [28]. In sum, the dual regulatory targeting of mitochondrial and chloroplast cistron expression by mTERF proteins to promote optimal energy production and oxygen consumption farther advances the evolutionary importance of OGE processes.
Conclusions
It is at present established that the same fix of functional genes are encoded in both the plastid and mitochondrial genomes, which limited the same conserved proteins in the electron transport concatenation [38]. Thus, it is strongly implied that OGE processes are critically linked to shared stereo-selective biochemical pathways. Maier and colleagues refer to this equally an case of parallel and convergent evolution. The ongoing processes underlying biologically meaningful evolutionary modification of the organellar genome tin exist partly attributed to regulatory stability of intracellular redox processes. As such, a hypothesis of evolutionary modification of intracellular redox regulation predicts that there is a specific location for the plastids and mitochondria genes that encode for bioenergetics membrane proteins that are functionally related to respiration or photosynthesis [38]. The dual evolution of the plastid and mitochondria genomes will effectively drive the retentiveness of functionally similar sets of ribosomal protein genes which are functionally required for proper ribosomal assembly.
It has been recently proposed that archaebacterium and eubacterium precursors led to the origin of eukaryotes [39,40]. Conversely, mitochondria arose from an alpha-proteobacterium and a eukaryote [40,41]. Plastids arose in a like style but from cyanobacterium and a eukaryote [forty]. Hence the eukaryotic cell was "adult". The developmental primacy of photosynthesis was probably due to abundant sunlight and coincident advent of requisite photovoltaic chemical processes. Furthermore, the byproducts of these processes, i.e., glucose and oxygen, introduced a major change in the biosphere with the associated evolutionary development of circuitous cellular respiratory processes and with major potential issues involving oxygen toxicity. In light of these changes, both photosynthetic and respiratory processes were driven by the potential for bacteria to farther enhance the intracellular membrane microdomains segregated according to functional physiological criteria.
Accordingly, the respiratory "bacterium" evolved and remained in place considering of its existential brokerage of molecular oxygen and the use of glucose every bit an initial fuel source in the bioenergetics of ATP production. In this regard, photosynthetic priming events promoted evolutionary acceleration of intracellular membrane differentiation, selective for plastid-like structures. This major contention is supported by the observation that many organelles tin can exist found in both plant and animal cells and that their molecular biology/bioenergetics share basic chemic processes.
The dual expression of mitochondria and functional chloroplasts within specialized creature cells indicates a high caste of biochemical identity, stereoselectivity, and conformational matching that are the probable keys to their functional presence and essential endosymbiotic activities for over ii.5 billion years [iii,42–44]. Thus, conformational matching imposes a loftier degree of rigidity on the systems, assuasive for their retention in development. Another component of the conformational matching hypothesis is that this phenomenon besides occurs via a chemical messenger and its receptor with the added fact that both must exist expressed simultaneously and accordingly on the right target [3,42–44]. Therefore, all the conformational dependent substrates and enzymes impose a rigidity on change in general, which does not favor change. Withal, change can and does occur considering slight changes may exist tolerated, giving rise to modified systems, east. g., the catecholamine pathway.
Footnotes
Conflict of interests
The authors declare no conflict of interests.
Source of back up: The report was, in part, funded by MitoGenetics, LLC (Sioux Falls, South Dakota)
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