Do Animal And Plant Cells Have Chloroplast
Med Sci Monit. 2015; 21: 2073–2078.
Mitochondria, Chloroplasts in Fauna and Plant Cells: Significance of Conformational Matching
Received 2015 May 25; Accepted 2015 Jun 26.
Abstract
Many commonalities between chloroplasts and mitochondria exist, thereby suggesting a common origin via a bacterial antecedent capable of enhanced ATP-dependent energy production functionally linked to cellular respiration and photosynthesis. Appropriately, the molecular evolution/memory 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 machinery for ATP synthesis in cellular respiration and photosynthesis. Furthermore, the dual regulatory targeting of mitochondrial and chloroplast cistron expression by mitochondrial transcription termination gene (MTERF) proteins to promote optimal energy production and oxygen consumption further advances these evolutionary contentions. As a functional issue of enhanced oxygen utilization and product, significant levels of reactive oxygen species (ROS) may be generated inside mitochondria and chloroplasts, which may effectively compromise cellular free energy production following prolonged stress/inflammationary weather condition. 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 observation of internalized algae within embryonic tissues of the spotted salamander strongly suggest that developmental processes within a vertebrate organism may require photosynthetic endosymbiosis as 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 ii.5 billion years.
MeSH Keywords: Chloroplasts, Kleptoplasty, Mitochondria, MTERF, PEWY, Reactive Oxygen Species, Stereospecificity
Background
Mitochondria and chloroplasts represent endosymbiont models of complex organelle development driven past evolutionary modification of permanently enslaved primordial leaner[ane–4]. Over various eukaryotic phyla mitochondria and chloroplasts either lone or together provide a concerted amplification of cellular energy production via shared biochemical pathways. Cellular dysregulation of these two distinct organelles may generate potentially dangerous reactive oxygen species (ROS) due to compromised complex bioenergetics energy product, systemic oxidative stress and compounded pro-inflammatory processes. Importantly, genetically- or biochemically-mediated failure of mitochondrial function in human populations represents a potentially dire factor in the etiology of major illness states that include Type Ii diabetes, atherosclerosis, rheumatoid arthritis, Alzheimer's Disease, and cancer progression [5–21]. In sum, these compelling mechanistic and clinical data advise that the extent of mitochondrial/chloroplast regulatory signaling may vary over the lifetime of the eukaryotic cell co-ordinate 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.e., a facultative anaerobic microbe with unlimited replication potential [24]. For example, anaerobic mitochondria in gill cilia of M. edulis have evolved to utilize the phenotype of a facultative anaerobe, demonstrating that this primitive type of respiration has been evolutionarily conserved [25,26]. Appropriately, anaerobically performance mitochondria may represent a re-emergence or evolutionary retrofit of primordial metabolic processes.
It has become recently apparent that mitochondria take discrete microenvironments composed of complex intracellular membrane structures with distinct functional identities determined by segregated biochemical pathways [27] (Figure 1). Given the shared chemical messengers between the two and interrelationships between the common energy processes it is non surprising that boosted commonalities are emerging. Furthermore, it is no surprise that mitochondria are present in both plants and animals, implying major shared regulatory, bioenergetic, and chemical substrate pathways. Commonalities of free energy processing in both plants and animals take become even stronger by the finding that chloroplast can be institute in animal cells. The discovery of kleptoplasty, a functional chloroplast in cells of a non-photosynthetic host [28] is a remarkable miracle [28–31]. It is too plant in metazoans, i.eastward., the sacoglossan body of water slug. Of equal importance is the longevity of functional kleptoplasts in the host, suggesting once more the mutual significance of bidirectional communication and the many commonalities in molecules exist so that this phenomenon can accept place and work. These bounding main slugs extract and incorporate functional chloroplasts from Ulvophyceae into their gut cells [32], assuasive their derived "nutrient" to be gained for months. The dependence on specific algae strongly suggests mutual bidirectional advice is responsible for these phenomena.
The prokaryotic cell is characterized past a general lack of highly structured intracellular organelles only displays intracellular regions of functionality with some membrane enhancements, e.g., mesosome. We surmise that with time this relatively elementary construction became more elaborate, adding membrane surface surface area to perform piece of work, enhancing a major part like respiration. In all probability the stimulus was solar free energy, causing the photolysis of water. This cell was driven in this direction considering information technology provided a new coping strategy for, counter-intuitively, Dna advancement. This evolving cellular architecture could not survive on its ain given the presence of by products it produced, due east.yard., ROS, which are basically toxic to unprotected intracellular components, notably Dna. This evolutionary self protection machinery was further advanced when oxygen levels increased as a consequence of photosynthesis. In all probability the cellular oxygen toxicity event was partly solved past having a "bacterium" develop in a "bacterium", becoming a eukaryotic cell, which could harvest specific bond free 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 free radical creator was effectively removed via chloroplasts, which originated in a like fashion as mitochondria. Thus, it is non surprising to detect both types of "leaner" in the same cell and others where only one is nowadays. Furthermore, given this close evolvement, enslavement was not an issue in this circumstance because each "cell" used the same or similar chemic messengers, stabilizing what appears to be a precarious relationship. Indeed, bidirectional communication served as the procedure for eukaryotic cellular communication/cooperation, which immune for metazoan evolution. Interestingly, metazoan evolution is still highly dependent on the intracellular communication with its endogenous bacterial components from which it evolved, east.g., intracellular and extracellular (gut microbiome). The vulnerability expresses itself in "mitochondrial dysfunction" in that it can exist so complicated and diverse depending on the tissue region affected. We further surmise that hypoxia plays a major role in triggering mitochondrial dysfunction since this entire relationship depends on a continuously ongoing energy processing system[2,21]. Briefly, the evolutionary advancement of eukaryotic cells requires this homeostatic free energy balance to maintain its multicomponent and faceted beingness. Any departure from the thermodynamically stabilized life course creates a pathology wherever information technology occurs. This process may also represent the deleterious mechanisms that may be associated with aging.
The ability of a chloroplast to office as a symbiotic bioenergetic organelle within the intracellular milieu of a representative invertebrate, i.e., the Sacoglossan ocean slug, was previously identified every bit a unique phenomenon unlikely to occur in vertebrates [28–32]. Recently, the observation of internalized algae within embryonic tissues of the spotted salamander strongly suggest that developmental processes inside a vertebrate organism may require photosynthetic endosymbiosis as an internal regulator [33]. Appropriately, information technology appears that dark-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 degrade equally the larvae develop over time [33]. Although endosymbiotic algal cells go through deposition, the cells can also encyst on the inner capsule wall which is detected through 18s rDNA distension in the reproductive tracts of the adult salamanders, thereby allowing for the transfer of genes from one generation to the next [33]. Due to the dense accumulation of algae within the embryo, a distinct light-green colour is exhibited which leads to beneficial effects for the embryo. Requisite physiological effects include lowering embryonic mortality, larger embryo size, and earlier hatching times. It is all the same unclear if the algae and the embryo have a true bidirectional symbiotic relationship because there is evidence that the algae take no increment in oxygen levels, simply they may benefit from the embryos when their nitrogenous waste material is released. In any event, this phenomenon defines a distinctive relationship between 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 every bit the PEWY motif in mitochondrial complexes, cyt b displays iv tetrapeptide residues (PDWY, PPWF, PVWY and PEWY) employed in catalytic reactions, which is at present identified as the Qo motif. PEWY, which is present in chloroplasts and mitochondria, and PDWY which is nowadays 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 development that the cyt bc1 complex with PEWY being the Qo motif will part all-time with a high potential and ubiquinone every bit its substrate [34]. For PDWY every bit the cyt b circuitous, a low potential and menaquinone will function the best. In sum, the molecular evolution/retentivity of the catalytic Qo quinol oxidation site of cytochrome b complexes, functionally underlies the common retention of a chemiosmotic proton slope mechanism for ATP synthesis in cellular respiration and photosynthesis.
The relationship betwixt photosynthesis and respiration tin vary, thereby demonstrating their dynamic nature. For example, when lycopersicon esculentum fruit ripen, their chloroplasts will change into photosynthetically inactive chromoplasts that can produce ATP through a respiration process known every bit chromorespiration [35]. Oxygen consumption through chromorespiration can be 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 6 f complex inhibitor 2,5-dibromo-3-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 exist acting as electron acceptors for the cytochrome b half dozen f circuitous. During ripening, mitochondrial numbers significantly decrease in the fruit tissue [35]. In lodge to compensate for this strong decrease, the number of chromoplasts will functionally increase during the later stages of ripening, thereby demonstrating critical modification of free energy processing.
Chiefly, plants require imported oxygen to bear out nigh of their biochemical reactions such as respiration fifty-fifty though they lack the power to distribute oxygen to the cells [36]. To compensate for the lack of this distribution mechanism, plants ofttimes display steep oxygen gradients that may be dumb due to environmental 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. As a key case, 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 Grouping-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, equally regulated by the Due north-end rule signaling pathway due to the impacted loss. The Northward-end rule signaling pathway represents a cellular response mechanism that requires ubiquitin ligation linked to proteasomal degradation via covalent modification of N-concluding amino acids [36].
Finally, the array of complex control mechanism past which organellar gene expression (OGE) promotes respiration, photosynthesis and plant development is actively under investigation [37]. Presently, several required components have been identified that have been functionally associated with OGE processes. Nucleus-encoded proteins take important roles in OGE by promoting various required functions such equally splicing, transcription, RNA processing and regulation of translational processes. Normative OGE is regulated by the family of mitochondrial transcription termination factors (mTERF). Mammalian mTERFS were originally proposed to specifically finish transcription, but further biochemical and molecular studies indicate that 3 out of the four mTERFS possess important regulatory activities necessary for ribosomal biogenesis and antisense transcription termination. Approximately 30 members of the mTERF family have been identified throughout plant development, but all the same little is known about how photosynthetic organisms are using mTERFs and OGE [28]. In sum, the dual regulatory targeting of mitochondrial and chloroplast factor expression by mTERF proteins to promote optimal free energy production and oxygen consumption further advances the evolutionary importance of OGE processes.
Conclusions
It is now established that the same gear up of functional genes are encoded in both the plastid and mitochondrial genomes, which limited the same conserved proteins in the electron transport chain [38]. Thus, it is strongly implied that OGE processes are critically linked to shared stereo-selective biochemical pathways. Maier and colleagues refer to this as an instance of parallel and convergent evolution. The ongoing processes underlying biologically meaningful evolutionary modification of the organellar genome tin can exist partly attributed to regulatory stability of intracellular redox processes. Equally 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 development of the plastid and mitochondria genomes will finer drive the retentiveness of functionally similar sets of ribosomal protein genes which are functionally required for proper ribosomal associates.
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 similar style but from cyanobacterium and a eukaryote [xl]. Hence the eukaryotic cell was "developed". The developmental primacy of photosynthesis was probably due to abundant sunlight and ancillary appearance of requisite photovoltaic chemical processes. Furthermore, the byproducts of these processes, i.east., glucose and oxygen, introduced a major change in the biosphere with the associated evolutionary evolution of circuitous cellular respiratory processes and with major potential bug involving oxygen toxicity. In light of these changes, both photosynthetic and respiratory processes were driven by the potential for bacteria to further enhance the intracellular membrane microdomains segregated according to functional physiological criteria.
Appropriately, the respiratory "bacterium" evolved and remained in place because of its existential brokerage of molecular oxygen and the utilise of glucose as an initial fuel source in the bioenergetics of ATP production. In this regard, photosynthetic priming events promoted evolutionary dispatch of intracellular membrane differentiation, selective for plastid-similar structures. This major contention is supported by the ascertainment that many organelles can exist found in both plant and beast cells and that their molecular biology/bioenergetics share bones chemical processes.
The dual expression of mitochondria and functional chloroplasts inside specialized animal cells indicates a high degree of biochemical identity, stereoselectivity, and conformational matching that are the probable keys to their functional presence and essential endosymbiotic activities for over ii.five billion years [iii,42–44]. Thus, conformational matching imposes a high degree of rigidity on the systems, allowing for their retention in evolution. 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 be expressed simultaneously and appropriately on the right target [3,42–44]. Therefore, all the conformational dependent substrates and enzymes impose a rigidity on alter in full general, which does not favor alter. Even so, change can and does occur considering slight changes may be tolerated, giving rise to modified systems, e. grand., the catecholamine pathway.
Footnotes
Conflict of interests
The authors declare no conflict of interests.
Source of support: The report was, in part, funded by MitoGenetics, LLC (Sioux Falls, South Dakota)
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