text:
Summary of potential roles of growth factors in CLG.
\r\n\tOn the other hand, smoothed particle dynamics (SPH) is a Lagrangian approach of CFD, which does not use mesh grids. Micro-hydrodynamics of granular particles can be solved using discrete element method (DEM) or granular dynamics. Till date, there is not a unified computational approache to solve fluid and particles together.
\r\n\tThis book aims to provide the state-of-the-art advanced CDF fundamentals and applications for emerging engineering processes. All cutting-edge CFD topics are welcome, but two main themes include extreme applications of CFD and advanced coupling of Eulerian and Lagrangian approaches. They are, for example, unprecedentedly large-scale parallel CFD simulations, development of novel OpenFOAM solvers, and coupled simulations of CFD, SPH, and DEM.
Pneumonectomy, the surgical removal of a lung, elicits a number of anatomical changes within the thoracic cavity that augments the diffusion capacity of the remaining lung. Pneumonectomy directs the entire cardiac output into the remaining lung and creates an empty hemithorax that results in a shift of the mediastinum toward the vacated thoracic compartment. In a number of experimental animal models, pneumonectomy initiates compensatory, regenerative growth of the remaining lung tissue that restores normal mass, structure and function. This growth process, called compensatory lung growth (CLG), is qualitatively similar across species, but differs with gender, age and hormonal status. CLG involves unique structure-function interactions not seen in solid organs. Little is known about the regenerative potential of human lungs. Although CLG has been reported in children after major lung resection, CLG in adults rarely occurs and remains a significant challenge. Mechanical feedback between the lung and thorax constitutes a major signal that sustains both post-natal lung development as well as post-pneumonectomy CLG. After pneumonectomy, increased mechanical stress and strain on the remaining lung induce adaptive responses to augment oxygen transport, including 1) recruitment of alveolar-capillary reserves, 2) remodeling of existing tissue, and 3) regenerative growth of acinar tissue when strain exceeds a critical threshold. This chapter will discuss the clinical aspects of pneumonectomy and will primarily review cellular and molecular mechanisms of CLG via experimental pneumonectomy models, which offers powerful insights into regenerative organ growth.\n\t\t
Before the 1930s, all pneumonectomies in humans were fatal due primarily to complications such as hemorrhage and sepsis. Another major challenge was to perform lung surgery with an open pneumothorax. A significant step to solve this problem was taken in 1903 by Ferdinand Sauerbruch, who designed a negative pressure chamber that allowed a team of surgeons to operate within the open chest without collapse of the lung (Sauerbruch, 1953). Sauerbruch brought his machine to New York in 1908, where Willy Meyer modified it to work with positive and negative pressure. Meyer successfully performed pulmonary resections in dogs in 1909 using suture closure of the bronchus and individual vessel ligation (Meyer, 1909), but he did not attempt his technique in humans. Years later, Quinby and Morse used a modification of this technique in dogs and showed that after pneumonectomy, the hemithorax fills with fluid and the remaining lung will shift to this empty side (Quinby & Morse, 1911). After the experimental use of endotracheal delivery of oxygen and anesthetics (Meltzer & Auer, 1909), Howard Lilienthal performed the first thoracotomy under endotracheal anesthesia at Mount Sinai Hospital (Lilienthal, 1910). He also had the largest published series of lobectomies, which he considered dangerous, reaching a mortality rate of 70% if more than one lobe was removed (Lilienthal, 1922). The first successful case of total pneumonectomy was described in 1931 by RoudolphNissen in Berlin. The patient was a 12 year-old girl with trauma injury of the left chest (Naef, 1987). She recovered completely after two months, and Nissen was quoted saying that the occlusion of the pulmonary artery did not cause cardiopulmonary collapse as predicted 20 years earlier by Quinby and Morse. A year later, Cameron Haight was the first surgeon to perform a successful pneumonectomy in the west at the University of Michigan, USA (Haight, 1934). This time, a 13 year-old girl developed pneumonia in the left lung and subsequent pyopneumothorax. After a small bronchial fistula, she recovered 3.5 months later. On April 1933, Evarts Graham performed the first successful pneumonectomy for cancer disease in a 48 year-old patient with a squamous cell cancer of the left upper lobe bronchus that survived almost 30 years after surgery (Graham & Berck, 1933). In his paper, Clarence Crawford standardized the pneumonectomy technique used for many years, including the use of periscapular incision, individual vessel ligation, suture closure of the bronchus and a new rhythmic ventilatory technique (O\'Shaughnessy & Crawford, 1938). In 1950, the introduction of a new double lumen tube allowed the ventilatory exclusion of the operated side (Bjork & Carlens, 1950). By the 1970s, a new technique changed the surgical method with the design of surgical staplers for lung resection. The first successful thoracic surgery performed using video assisted thoracic surgery (VDATS) was for treatment of pneumothorax in 1990. Shortly thereafter the first descriptions of pulmonary lobectomies and pneumonectomies appeared (Davies & Panasuk, 1992). There has been steady progress in thoracic surgery and pneumonectomy in particular through the years, with advances in knowledge of respiratory physiology, anesthetics and ventilation techniques, as well as more sophisticated methods of lung resection. These advancements have transformed pneumonectomy surgery from a "dangerous procedure" to become a very useful treatment for both malignant and non malignant diseases of the lung and airways.\n\t\t\t
Pneumonectomy is known to be associated with high morbidity and mortality. However, in certain instances, it offers the only chance for a cure. The indications for pneumonectomy are usually classified in two major groups: pneumonectomy for benign disease and pneumonectomy for malignant disease. Due to the improvement of antimicrobial therapies and better control of inflammatory diseases, pneumonectomy for benign diseases is not a routine procedure in our times. The conditions considered in this group belong to several categories including inflammatory, traumatic, congenital and other miscellaneous conditions (Conlan & Kopec, 1999). These conditions carry a high mortality rate in most cases, but some examples of non-malignant disease have an excellent cure rate. Indications for pneumonectomy in cases of bacterial or fungal infections include symptomatic patients with hemoptysis, productive cough or chronic empyema, as well as unilateral lung destruction documented by CT or bronchography(Blyth, 2000). In patients with pulmonary tuberculosis, pneumonectomy is indicated for either multidrug resistance disease or for complications or sequelae of tuberculosis infection. The most common fungal infection that requires pneumonectomy is produced by Aspergillum, which can produce severe infections with recurrent or massive hemoptysis in over 75% of patients (Conlan, et al., 1987). Pneumonectomy for trauma is uncommon, but is associated with high morbidity and mortality (66-75%). The most obvious indication of the procedure is the laceration of the lung and airways, which can produce massive hemorrhage and air leakage. Complications of congenital and other miscellaneous lung diseases, as well as completion pneumonectomy are rare indications for pneumonectomy,which can be associated with high mortality. Pneumonectomy for malignant disease has become the most common indication for lung resection today, which includes both primary lung tumors and metastatic lung disease. It is regarded as the only curative treatment for non small cell lung cancer (Shields, 1982) and also as an effective therapeutic option for pulmonary metastasis; however, these can be associated with high mortality (Spaggiari, et al., 1998).\n\t\t\t
Pneumonectomy is associated with a 38-59% rate of morbidity and a 30-day mortality ranging from 3-12%. Postoperative cardiac dysrhythmias (e.g., atrial fibrillation and supraventricular tachycardia) are relatively common complications occurring in approximately 20-40% of patients following pneumonectomy. Postpneumonectomy pulmonary edema (PPE) and acute respiratory distress syndrome (ARDS) occur in 4-7% of patients and are increasingly believed to be the same disease process. PPE/ARDS results in noncardiogenic pulmonary edema and is manifest by diffuse pulmonary infiltrates on chest radiograph combined with profound hypoxia and respiratory failure frequently requiring mechanical ventilation. One of the more devastating complications of pneumonectomy is an empyema involving the postpneumonectomy space. A postpneumonectomy empyema is usually associated with a bronchopleural fistula (BPF), which is a communication between the mainstem bronchial stump and the pleural cavity. The incidence of BPF and empyema ranges from 2-8%, but both complications are significantly more common in patients who undergo pneumonectomy for septic pulmonary disease (e.g., tuberculosis or fungal disease) (Deschamps, et al., 2001). Postpneumonectomy syndrome is a rare complication characterized by stridor, dyspnea and recurring pneumonia. Postpneumonectomy syndrome is more common following right pneumonectomy and results from severe shifting of the mediastinum and contralateral lung into the postpneumonectomy pleural space, which in turn leads to compression of the contralateral mainstem bronchus between the vertebral bodies and descending aorta (Kopec, et al., 1998).\n\t\t\t
Pneumonectomy results in reduced lung function. Although residual volume (RV) declines after pneumonectomy, it decreases less than expected as a result of the hyperexpansion that occurs in the remaining lung. Forced expiratory volume in one second (FEV1) and forced vital capacity (FVC) both typically decrease by 25-40%. Diffusion capacity usually decreases by less than 50% whereas PaO2 and PaCO2 typically remain unchanged from preoperative levels (Kopec, et al., 1998).\n\t\t\t
The removal of a lung entails profound mechanical, metabolic and vascular changes in response to the reduction in lung volume and the empty space in the hemithorax. These changes trigger a compensatory responsein experimental models, known as CLG, which are directed toward reestablishing the normal rate of oxygen exchange capacity. This section describes a number of experimental animal models of CLG that are currently used to study this process, as well as a detailed evaluation of the CLG response in these models, including lung morphometry and imaging.\n\t\t\t
In many animal models, pneumonectomy (or lobectomy) induces CLG of the remaining lung, resulting in rapid restoration of total lung volume, compliance, mass, DNA, protein, alveolar number, and normal lung cell populations. Pneumonectomy has proven to be a reliable model for characterizing the sources, mechanisms, and functional limits of the compensatory growth response after removal of lung tissue. Pulmonary resections in animals began in 1881 when it was documented that the remaining lung eventually expands to the same size as both lungs. Early animal studies established the basis for application of the procedure to humans, beginning in the 1900s. Cohn in 1939 first established mechanical forces as a major signal for the compensatory increase in lung mass following lobectomy. In the 1950s, Schilling detailed the well-preserved functional status in dogs that underwent removal in stages of up to nearly 70% of lung mass (Schilling, et al., 1958). The use of small animals (e.g. rabbit, rat, and mouse) from the 1960s to the current day has had a great impact in uncovering the hormonal, cellular, and molecular responses to pneumonectomy(Bennett, et al., 1985; Buhain & Brody, 1973; Rannels, et al., 1979; Romanova, et al., 1967). Significant progress in understanding the cellular and molecular pathways of tissue regeneration in vertebrates were achieved using both transgenic mice and molecular biology techniques (Leuwerke, et al., 2002; Sakamaki, et al., 2002; Sakuma, et al., 2002). Functional compensation to pneumonectomy has been described mainly in dogs to define the limits of such compensation (Ravikumar, et al., 2004; Takeda, et al., 1997).\n\t\t\t
Alveolar epithelial cells in pneumonectomized rats exhibit metabolic changes typical of accelerated cell growth. Studies in mice and rats indicate that type II epithelial cell hypertrophy, proliferation and differentiation into type I cells characterize CLG in a fashion similar to early postnatal lung growth and lung repair after injury (\n\t\t\t\t\t\tKaza, et al., 2002\n\t\t\t\t\t).\n\t\t\t\t
The post-pneumonectomy CLG response is independent of the lobe or lobes removed in small animals; and all remaining lobes grow rapidly until normal total lung mass is restored. Increases in lung volume after pneumonectomy parallel accumulation of tissue growth. However, the increase of growth in the remaining lobes is not uniform (Fernandez, et al., 2007). Development of sophisticated morphometric methods has permitted accurate analysis of lung volume and alveolar number, and studies indicate that new alveoli are formed during post-pneumonectomy CLG (\n\t\t\t\t\t\tKaza, et al., 2002\n\t\t\t\t\t; Sakamaki, et al., 2002). In dogs, however, it appears that the CLG response is initiated after a certain threshold is achieved (removal of >50% of total lung mass) (Hsia, et al., 1994). Bronchoalveolar stem cells (BASC) have also been implicated in CLG. Their proliferation and differentiation into alveolar epithelial cells type II and I, contribute between 0-25% to the regenerative lung growth process (Nolen-Walston, et al., 2008).\n\t\t\t
Vascular growth and angiogenesis during CLG has not been well characterized. Stimuli known to initiate angiogenesis include hypoxia, inflammation, and mechanical factors such as shear stress and stretch. Our laboratory has shownthat angiogenesis is necessary for successful CLG by demonstrating that angiogenesis inhibitors such as fumagillin or thalidomide prevented increased lung weight and volume after pneumonectomy(Maxey, et al., 2003). We have also shown that pneumonectomy induces arterial growth including the increase in length and number of branches of pulmonary arteries and that these changes are proportional to the amount of tissue removed (Le Cras, et al., 2005). When a bilobectomy was performed in rats (24% of lung tissue removed), the arterial area of the remaining lung increased by 26% compared to sham animals. Furthermore, when lung resection was more extensive (trilobectomy, 52%) we found that the increase in arterial area increased by 47% (Le Cras, et al., 2005). Other researchers have shown the effects of exogenous angiogenic factors, such as vascular endothelial growth factor (VEGF) in the mouse model. Additional VEGF therapy accelerated the CLG response, which was completed in only 4 days compared to 10 days in the pneumonectomy control group (Sakurai, et al., 2007).\n\t\t\t
Several general hypotheses have been advanced to account for events that initiate the cellular and molecular changes that lead to CLG. Mechanical signals, transient hypoxia associated with thoracotomy, and elevated blood flow have been considered; however, no single event has been proven to account for the growth response.\n\t\t\t
After resection of the lung, increased inflation of the remaining lung and increased blood flow will induce stretch in both alveolar and endothelial structures. The displacement of the lung to the empty hemithorax is also a feature of CLG. These mechanical forces have an important role in initiating and regulating CLG as it was demonstrated when lateral displacement of the lung was restricted by the use of an inflatable prosthesis. This prevention of the mediastinal shift significantly limited mechanical lung strain, and CLG was thus significantly impaired by30-60% (Hsia, et al., 2001). Another factor, increased alveolar inflation, was also implicated when experiments using in vitro perfusion of lungs with or without constant positive pressure ventilation (CPAP) of 20 cmH2O, demonstrated that lungs with increased inflation had cellular hyperplastic changes, such as elevated levels of cAMP and PKA activity, but perfusion alone did not account for these changes (Russo, et al., 1989). It is clear that CLG is very complex with multiple metabolic factors. Hyperinflation and stretch applied to the remaining lung after pneumonectomy are powerful signals to initiate CLG, and it is known that stretch of alveolar cells induces important changes associated with cell growth and septal formation, including signal transduction, protein turnover, growth factor production, proliferation, and apoptosis (Brody, 1975; Davies, et al., 1982; Fehrenbach, et al., 2008; Karl, et al., 1989). It appears that lung stress and strain generated after pneumonectomy, overlaid on a background of heightened developmental lung strain generated by the expanding thorax, intensifies the CLG responses. Importantly, minimizing post-pneumonectomy strain of the remaining lung with space-occupying, inert material blunts the CLG response.\n\t\t\t
Post-pneumonectomy changes in pulmonary blood flow have been considered as possible signals for CLG. Increased perfusion, reflecting elevated cardiac output to the remaining lung, likely causes physical distention of the pulmonary vasculature, resulting in a mechanical signal for lung growth and a concurrent increase in the growth factor and/or nutrient availability to the lung. It has been described that, after banding of the left caudal pulmonary artery in ferrets that reduced blood flow to the lung by 25%, CLG still occurred after pneumonectomy. However, the caudal lobes in the banded animals were 17% smaller than those of non-banded animals and tended to have lower protein content (McBride, et al., 1992). In our laboratory, we have shown that after left pneumonectomy, increases in growth and proliferation were not uniform among the right lobes but were greater in the upper and cardiac lobes. These unequal changes coincided with a predominant vascular growth in the upper lobe, which received the highest fraction of relative blood flow (Fernandez, et al., 2007).\n\t\t\t
Hypoxia has been shown to stimulate alveolar growth either directly or via interaction with other signals. The effects of hypoxia after pneumonectomy were initially described in the rat model, where pneumonectomized rats that recovered at hypoxic levels showed significant increases in lung weight and volume indices, increased alveolar surface area and total alveolar numbers compared to normoxic and hyperoxic rats (Sekhon, et al., 1993). Hypoxia-inducible factors (HIFs), which are activated in response to oxidative stress, hypoxia, injury, and physical forces, regulate transcription of genes involved in a wide array of functions including glycolysis, erythropoiesis, apoptosis, and angiogenesis. Most of these stressors are directly or indirectly associated with a change of intracellular oxygen tension, which leads to stabilization of the HIF-1α protein and increases the transcriptional activation of target genes. Elevated hypoxia-induced mitogenic factor (HIMF) and HIF-1α mRNA and protein expression has been documented during CLG, thus these pathways may play an important role in mediating CLG (Li, et al., 2005; Zhang, et al., 2006; Zhang, et al., 2007).\n\t\t\t
The molecular mediators of CLG remain poorly understood. CLG involves regulated pathways of cell cycle activity, cell differentiation, synthesis and organization of connective tissue components, tissue remodeling, and angiogenesis. Studies in animals have led to several hypotheses that various pathways play a role in the induction of CLG including post-operative release of hormones and growth factors, as well as changes in cellular behavior. This section describes several important aspects of post-pneumonectomyCLG regulatory mechanisms.\n\t\t\t
The most substantial evidence for hormonal regulation of CLG stems from investigations that involved surgical ablation of the adrenal glands, or adrenalectomy, which alone does not stimulate lung growth. Adrenalectomy performed prior to pneumonectomy increases the rate and extent of CLG above that observed in rats after pneumonectomy alone. This stimulatory effect on lung growth was blocked by daily doses of hydrocortisone acetate, evidenced by parameters such as dry lung weigh and DNA content, which were similar to the pneumonectomy group (Bennett, et al., 1985). This blocking effect was found only if the therapy was used continuously for the entire period after surgery (Rannels, et al., 1987). A combination of dexamethasone, 8-bromo-3\'-5\'-cAMP and isobutylmethyl-xanthine (DCI) has been successfully used to accelerate CLG in mice, represented by an increase in lung dry weight index and an increased number of alveoli by morphometric analysis. The effect of a single airway dose was enough to maintain the effect for the entire 28-day period of study. This effect seems to be modulated by thyroid transcription factor 1 (TTF-1), since its transient inhibition attenuated CLG (Takahashi, et al., 2011). Adrenal glucocorticosteroids seem to have a role in the modulation of the accelerated CLG initiated by pneumonectomy. Several lines of evidence suggest a possible role of growth hormone in the regulation of CLG. Significantly higher serum levels of growth hormone were detected in pregnant rats 3 days after pneumonectomy when compared with sham and unoperated rats (Khadempour, et al., 1992). In diabetic rats, which normally have greater levels of growth hormone and adrenal corticosteroids, pneumonectomy was accompanied by an increased dry lung weight index as well as higher elastin and collagen content, when compared to control pneumonectomy and sham rats (Ofulue & Thurlbeck, 1995). Following pneumonectomy, rats implanted with a subcutaneous growth hormone-secreting tumor (MtTF4) underwent a CLG response similar to non-tumor-bearing controls; however, lung growth in MtTF4 rats was associated with a greater lung volume.\n\t\t\t
There is ample evidence that growth factors regulate CLG, and the production of many growth factors is known to be sensitive to mechanical strain. Each growth factor modulates different aspects of cellular growth, but any one growth factor cannot recapitulate the entire CLG response, and there is much functional overlap among growth factors. Our laboratory, among others, has demonstrated important roles for epidermal growth factor (EGF), hypoxia-induced mitogenic factor (HIMF), keratinocyte growth factor (KGF) and retinoic acid in CLG. Other growth factor signaling pathways have been found to be activated after PNX including insulin-like growth factor-1 (IGF-1), hepatocyte growth factor (HGF), erythropoietin receptor, and hypoxia-inducible factor-1 (HIF-1α). These growth factors will be discussed below.\n\t\t\t\t
It is been shown that EGF via its receptor (EGFR) plays a role in prenatal and postnatal lung development. Its actions involve the synthesis of surfactant precursors and the differentiation of type 2 epithelial cells. Using a pig lobectomy model, our laboratory demonstrated an upregulation of EGFR expression two weeks after lobectomy, which coincided with an increased alveolar cell proliferation index of the remaining lung. At 3 months after surgery, there was an increase in the lung protein/DNA ratio in the lobectomy group compared to controls (\n\t\t\t\t\t\t\tKaza, et al., 2001\n\t\t\t\t\t\t; Kaza, et al., 2002). We also have documented the effects of EGF in CLG using a rodent pneumonectomy model. When exogenous EGF was administered to rats after pneumonectomy, it caused significantly higher lung weight and volume indices when compared to pneumonectomy control animals. We also detected an upregulation of EGFR after exogenous EGF supplementation (Kaza, et al., 2000), suggesting that the upregulation of EGF signaling is a feature of this process and is capable of modulating post-pneumonectomy GLG.\n\t\t\t\t
Erythropoietin (EPO) actions have been classically associated with the induction of erythropoiesis; however, organ specific EPO receptor (EPOR) signaling is also involved in development, angiogenesis and organ growth. Researchers have shown that EPOR is upregulated both during postnatal lung maturation and during CLG in adult dogs (Foster, et al., 2004). Furthermore, they demonstrated an upregulation of one of its upstream activators, HIF-1α, during the same processes. Using an in vitro system, the same group showed that upregulation of HIF-1αin cultured HEK-293 cells also caused the upregulation of endogenous EPOR expression (Zhang, et al., 2006). This also provided evidence of a possible role of EPOR in CLG after pneumonectomy.\n\t\t\t\t
HGF is known to selectively stimulate epithelial and endothelial cells, and the major sources for HGF in the lung are macrophages, fibroblasts, and endothelial cells. The increase of serum HGF during the first week after major lung resections in humans has been documented (Dikmen, et al., 2006; Sugahara, et al., 1998). A more comprehensive study of the role of HGF in CLG was performed using a mouse pneumonectomy model (Sakamaki, et al., 2002). In this study, the authors demonstrated an increased level of both lung mRNA and protein expression of HGF after pneumonectomy, and serum HGF levels were also higher when compared to sham operated animals. These findings were accompanied by an increased proliferation index of alveolar and airway epithelial cells, which peaked at day 5 after surgery. They also detected an upregulation of the HGF receptor (c-Met) at day 3 post-pneumonectomy. In addition, injections of HGF twice daily enhanced the proliferative response of these cells as well as increased lung weight index at day 3 when compared to controls. Use of a neutralizing antibody against HGF resulted in the inhibition, although incomplete, of the increase in lung weight and DNA synthesis observed. Another interesting study evaluated the effects of CLG in a rat model of elastase-induced emphysema, with transfection of the human HGF cDNA into the lung (Shigemura, et al., 2005) or implantation of adipose tissue-derived stromal cells (ASCs), which produce a large amount of angiogenic factors including HGF (Shigemura, et al., 2006). Therapy with gene transfection was performed using the hemagglutinating virus of Japan (HVJ) envelope-plasmid complex. In the HGF- and ASC-treated animals, increased levels of both exogenous and endogenous HGF were detected; and furthermore, HGF enhanced the CLG response by increasing lung cell proliferation and improving functional parameters. Taken together, these studies provide strong evidence for a role of HGF in the proliferative responses during CLG.\n\t\t\t\t
In a collaborative study, we demonstrated the role of HIMF in the context of CLG (Li, et al., 2005). The mRNA and protein expression of HIMF, which is known by its mitogenic and angiogenic properties, was upregulated after pneumonectomy (days 3-14) when compared to sham operated mice. The elevated HIMF expression also coincided with an increase in cell proliferation index in lungs of these animals. HIMF expression after pneumonectomy was mainly detected in airway epithelial, endothelial and type 2 epithelial cells. Intratracheal instillation of exogenous HIMF increased the proliferative activity in these cells, documenting its mitogenic properties and establishing its role in CLG.\n\t\t\t\t
Researchers have shown that HIF-1α expression is upregulated both during postnatal lung maturation and during adult CLG in dogs and that this coincides with the upregulation of one of its downstream targets, EPOR. In vitro experiments also provided evidence that upregulation of HIF-1αin cultured HEK-293 cells triggers the upregulation of endogenous EPOR expression (Zhang, et al., 2006). Another study found that lung expansion is a major contributor to the activation and stabilization of HIF-1α. Acute deflation of prosthesis in the chest cavity of pneumonectomized dogs triggered the increase of both HIF-1 and several HIF-1 targets including EPOR and VEGF compared to non-deflated animals. They concluded that these increases did not depend on hypoxia but instead were due to stretch-related signals after lung resection (Zhang, et al., 2007).\n\t\t\t\t
IGF-1, its receptor and binding proteins are naturally expressed in the lung during development, and IGF-1 is known to contribute to the regulation of postnatal lung cell proliferation. Researchers have shown that 2 and 6 days after pneumonectomy in rats, the bronchoalveolar lavage fluid from these animals demonstrated significantly increased mitogenic activity when applied in vitro to fibroblasts compared with controls. Importantly, such activity was partially inhibited by the use of neutralizing antibody against IGF-1, and the levels of IGF-1 were elevated by 100% at day 2 after pneumonectomy(McAnulty, et al., 1992). In a separate study, IGF-1 mRNA expression was again significantly increased after 21 days post-pneumonectomy in a model of neonatal CLG in lambs (Nobuhara, et al., 1998).\n\t\t\t\t
KGF has been shown to play an important role in alveolar epithelial cell proliferation and lung development. In our laboratory, exogenous KGF administered to rats after pneumonectomy further enhanced several parameters of CLG compared to control animals. Changes in lung weight index, lung volume index as well as morphometric parameters were accompanied by a significant increase in pulmonary cell proliferation index, providing the first evidence for a role of KGF in CLG (\n\t\t\t\t\t\t\tKaza, et al., 2002\n\t\t\t\t\t\t). A more recent study corroborated our findings where epithelial cell proliferation was further enhanced after in vivo transfection of a KGF cDNA vector in a model of CLG in rats, confirming KGF as an important lung mitogenic factor (Matsumoto, et al., 2009).\n\t\t\t\t
Retinoic acid, a metabolite of vitamin A, has been implicated in normal lung development and cell proliferation. Our laboratory described the effects of exogenous retinoic acid during CLG in a rat model (\n\t\t\t\t\t\t\tKaza, et al., 2001\n\t\t\t\t\t\t). At 10 and 21 days after pneumonectomy, lung weight, lung volume and cellular proliferation indices were all significantly augmented in rats that received exogenous retinoic acid versus vehicle controls. Interestingly, we also found that pulmonary expression of EGFR was upregulated in lungs after retinoic acid treatment, uncovering a possible relationship between these two important growth factors in CLG. Another similarly designed study corroborated our results several years later (Karapolat, et al., 2008); however, these effects did not translate into functional recovery according to studies developed using the dog model (Dane, et al., 2004).\n\t\t\t\t
Angiogenesis, the formation of new blood vessels, is a critical step in normal organ development and in abnormal processes such as tumor growth and metastasis. In the lung, alveolar growth and angiogenesis should occur concurrently in order to result in normal organ development or regeneration. One of the most important angiogenic growth factors, VEGF has been studied in CLG, revealing its importance in this regenerative process. Researchers described the effects of exogenous VEGF in the mouse model, showing that exogenous VEGF therapy accelerated the CLG response, which was completed in only 4 days compared to 10 days in control animals. However, these effects could not be blocked by the use of either VEGF receptor inhibitors or neutralizing antibodies (Sakurai, et al., 2007). Another study associated lung expansion and other signaling pathways to VEGF. The acute deflation of prosthesis in the chest cavity of pneumonectomized dogs triggered the increase of HIF-1α and its targets EPOR and VEGF compared to non-deflated animals, showing the interaction between these signals in the regulation of CLG (Zhang, et al., 2007). A recent paper studied the modulation of different VEGF isoforms along with its receptor during CLG, where VEGF 188 mRNA expression was decreased compared to sham animals and VEGF 164 and VEGF 120 mRNAs increased during days 1 and 3 respectively, describing what they believe is the recapitulation of the pattern of expression for these isoforms in the fetal lung (Jancelewicz, et al., 2010). A brief summary of evidence for the role of growth factors in CLG is shown in Table 1.\n\t\t\t\t\t
|Growth Factor\n\t\t\t\t\t\t\t\t
|Pneumonectomy increased EGFR lung expression.\n\t\t\t\t\t\t\t\t
Exogenous EGF enhances CLG.
|Rat, pig\n\t\t\t\t\t\t\t
|Pneumonectomy increased EPOR lung expression.\n\t\t\t\t\t\t\t\t
|Pneumonectomy increased serum HGF, along with expression of HGF and its receptor (c-Met) in lung.\n\t\t\t\t\t\t\t\t
Exogenous HGF enhances CLG.
Neutralizing antibody against HGF attenuate CLG.
Mouse, rat
|Pneumonectomy increased lung HIMF mRNA and protein expression.\n\t\t\t\t\t\t\t\t
Exogenous HIMF enhances proliferation.
|Pneumonectomy increased lung HIF-1α mRNA and proteinexpression.\n\t\t\t\t\t\t\t\t
Lung expansion modulates HIF-1αexpression.
|Pneumonectomy increased lung IGF-1 mRNA and protein expression.\n\t\t\t\t\t\t\t\t
Neutralizing antibody against IGF-1 attenuate mitogenic in vitro effects.
|Lamb, rat\n\t\t\t\t\t\t\t
|Pneumonectomy increased lung KGF mRNA.\n\t\t\t\t\t\t\t\t
Exogenous KGF enhances CLG.
|Retinoic Acid\n\t\t\t\t\t\t\t\t
|Exogenous retinoic acid enhances CLG.\n\t\t\t\t\t\t\t\t
Exogenous retinoic acid induces EGFR.
|Pneumonectomy increased lung VEGF 164 and 120 mRNA expressions.\n\t\t\t\t\t\t\t\t
Exogenous VEGF accelerates CLG.
Pneumonectomy induces shear stress in the lung, and several key transcription factors provide links between shear-mediated signaling and CLG. Mitogen-activated protein kinases (MAPKs), composed of extracellular signal-regulated kinases (ERKs), c-jun NH2-terminal kinases (JNKs), and p38 MAPKs, play a critical role in cell differentiation, growth and apoptosis and the regulation of various transcription factors and gene expression. One of the first reports about stretch-induced early gene expression demonstrated that in as early as 30 minutes post-pneumonectomy in rats, c-fos and JunBare significantly increased. These results were also reproduced when in vitro ventilation-perfusion was used (Gilbert & Rannels, 1998). Using array technology, researchers have also shown significant increases in six different transcription factors as early as 6 hours after pneumonectomy in mice including Erg-1 and Nur77, all of which have important roles in vascular biology, development and stress response (Landesberg, et al., 2001).In addition, a more recent paper found an important role for thyroid transcription factor 1 (TTF-1) in CLG (Takahashi, et al., 2010). These investigators detected a significant increase of TTF-1 expression 12 hours after pneumonectomy with TTF-1 expression primarily observed in cells of the alveolar ducts. When TTF-1 was repressed using small inhibitory RNAs (siRNAs), the CLG response was also temporally delayed.\n\t\t\t
Telomerase is an important enzyme for DNA repair and contributes to cell maintenance. Telomerase prevents the excess shortening of telomeres, and it has been demonstrated that telomerase is active in proliferating cells in a number of organs, including the lung. In humans, mutations of the telomerase gene results in a pathological condition known as idiopathic pulmonary fibrosis (IPF). Its role in preserving lung epithelial integrity was demonstrated in an experimental model in mice, where telomerase deficiency resulted in a reduction in the number and integrity of type 2 alveolar epithelial cells (AEC2) (Lee, et al., 2009). It is important to know that this defect may not be apparent in early generations, but after progressive inbreeding, it was possible to establish the deficiency in AEC2s as well as in bronchoalveolar stem cell (BASC) populations in the lung, due to shortening of the telomeres. In a recent study, the importance of telomerase was demonstrated for CLG after pneumonectomy using telomerase deficient mice from second (F2), third (F3) and fourth (F4) generation animals (Jackson, et al., 2011). In wild-type mice, the activity of telomerase was found to increase in isolated AEC2s up to 3.5-fold during post-pneumonectomy days 3 and 7. In addition, the total number of AEC2s and BASCs also increased at day 3 after pneumonectomy. However, pneumonectomy resulted in diminished CLG in F3 telomerase null animals, expressed as failure to induce an increase in lung mass by day 7 after pneumonectomy. In addition, the number of AEC2s and BASCs did not increase during the initial period after pneumonectomy when compared to wild-type mice. The normal CLG, both in lung mass and AEC2 numbers, in wild-type mice was also accompanied by an elevation in the proliferation marker Ki-67, early growth response gene (Egr-1) and repair transcription factors such as ERK 1/2, which were not observed in telomerase null mice. The authors concluded that telomerase deletion produces an attenuated CLG response after pneumonectomy by arresting cell growth and inducing DNA damage.\n\t\t\t
The extent of CLG in humans following pneumonectomy or lobectomy is incompletely investigated, but a number of long-term physiological studies suggest, however, that some degree of CLG may occur, especially in children (Nakajima, et al., 1998; Nonoyama, et al., 1986). The ability to manipulate the gain/loss of function for a particular gene in experimental animals has begun to provide a more detailed understanding of the molecular mediators and the pathologic consequences of CLG. Also, recent developments in cell therapy of diseased lungs with the use of adipose stem cells are indeed promising (Shigemura, et al., 2006). An important long-term goal of research into mechanisms of CLG is to generate knowledge that will allow the induction of alveolar regeneration or that rescues failed alveologenesis in humans. Such understanding will facilitate the development of therapies for the management of end-stage lung disease, lung volume reduction surgery, and transplantation. The potential clinical applications of this research are great. Specific examples of patients who would clearly benefit from lung regenerative therapies include chronic obstructive pulmonary disease (COPD), emphysema, pulmonary hypertension, bronchopulmonary dysplasia (BPD), as well as premature infants whose lungs are too underdeveloped to support life.\n\t\t
The objective of this chapter is to use a simple classical Mach-Zehnder interferometer separately under spatial-fringe-mode (Poynting vectors noncollinear in the output) and scanning-fringe-mode (Poynting vectors perfectly collinear in the output). The purpose is to demonstrate that "single photon interference" is a noncausal interpretation of the light-matter interaction process, where light energy absorbed by a detector array represents bright fringes and nonabsorption represents dark fringes. The process is not due to some mystical arrival and nonarrival of "photons" that defies the diffractive light propagation physics. The light propagation physics cannot remain strictly valid everywhere except in human-constructed interferometers and Young's double-slit diffractometers. We are using mathematical formalism, which we have accepted for a couple of centuries. We support our arguments based upon strict causality built into our mathematics and corroborated by measured data. This chapter considers light-matter interaction process exclusively from the standpoint of semi-classical model, where light is treated as classical electromagnetic waves and light detecting atoms and molecules as quantum entities that have discrete quantum mechanical binding energy levels (or bands) with finite transition probabilities.
In Sections 2.1 and 2.2, we use simple Mach-Zehnder interferometers with the Poynting vectors of the two combined output beams under (i) noncollinear and (ii) strict collinear conditions. The experiments demonstrate that the two incident signals from the two opposite sides of the beam combiner must be simultaneously present to generate the observed superposition effects. The simple mathematical formalism tells us that wave amplitudes by themselves never re-arrange their spatial or temporal energy distribution. They continue to propagate unperturbed by each other's physical presence. We have called this wave property, noninteraction of waves (NIW). In Section 3, we show a simple derivation of the Einstein's photoelectric equation as a superposition effect congruent with the semi-classical model. Since the basic mathematical formalism of Quantum Mechanics (QM) is correct, we must learn to avoid being trapped in the century-old and un-necessary circular arguments behind diverse mystical interpretation of Quantum Mechanics (QM). In Section 4, we describe this briefly by underscoring that the so-called "Measurement Problem" of QM is a fundamental "Information Retrieval Problem" out of all experiments, which cannot be solved by promoting elegant and beautiful mathematical theorems. It will require consistent and iterative attempts on the part of generations of scientists to visualize the invisible interaction processes going on in nature. Experimental data driven "evidence-based science" is the best approach to unravel nature's causal interaction processes while guided by strictly causal mathematical rules. Unfortunately, most of the details of the micro interaction processes are still beyond our capability to visualize directly. The problem is further compounded by the fact that the beautiful mathematical logics are inventions by the "subjective" neural networks of the human species.
The most neglected issue in current books and literature is that the co-propagating and cross-propagating wave amplitudes pass through each other completely unperturbed (uninfluenced) by each other's presence in the absence of interacting materials. In other words, the superposed wave fronts by themselves do not generate observable interferometric fringes whenever they are superposed. This is noninteraction of wave (NIW) amplitudes [1]. Alhazen observed this phenomenon almost 1000 years ago using a set of candles and a pinhole camera [2]. Huygens underscored this in his book [3] around 1667. This is why it is important to remember that even quantum electrodynamics acknowledges that photon-photon interaction cross section is immeasurably small [4]. In fact, Dirac mathematically found that "different photons do not interfere with each other" (NIW?). Unfortunately, he introduced the noncausal notion that "a photon interferes only with itself" [5]. This assertion is noncausal because interference fringes always appear as some physical transformation in a detector induced by more than one amplitude signals carrying different phase information. Further, the dark fringes are not due to nonarrival of "photons." It is because the joint stimulations by the out-of-phase E-vectors (only when equal amplitude!) fail to stimulate the detecting dipoles, and hence, the field energy cannot enter into the detecting dipoles' quantum cups. We always represent the superposition equation by two separate amplitude terms, each containing its own phase factor representing separate and independent oscillations of the E-vectors. A single stable elementary particle (here, a "photon") could not be multivalued in its critical dynamic parameters at any single moment. Further, in the absence of any other interacting force(s), it should not, by itself, appear or disappear in some specific physical location for some specific instrumental alignments made by humans to generate dark and bright fringes. In writing supposition equation, we always completely ignore the mathematics necessary to represent inherent diffraction physics of light waves. This is only partially correct for collimated beams traveling only short distances. Under this assumption, we neglect spatial evolution of complex amplitudes of the two waves. Photons cannot independently change their trajectories. Propagation of EM waves is rigorously given by Maxwell's wave equation and Huygens-Fresnel diffraction integral. Noninteraction of photons or generalized noninteraction of wave (NIW) [1] is equivalent to the statement that the linear mathematical superposition principle (SP) does not represent an observable phenomenon. It is a correct mathematical starting point to derive the observable superposition effect (SE) as the nonlinear square modulus of the expression for the linear superposition principle. In spite of this existing knowledge, somehow we get systematic training to accept that individual "indivisible light quanta" interfere by themselves. The phrase, "indivisible light quanta," represents energy "hν," a quantum cup of energy that is exchanged between quantized materials and classical waves. The mathematical expression "hν" is devoid of phase information. Note that our equation representing superposition phenomenon consists of two or more superposed waves with different amplitudes and phases carried by separate waves. Insertion of "bra" and "ket" symbols on to the complex amplitude terms fails to accommodate the physical properties behind the spontaneous and perpetually diffractive EM waves through any medium or instrument (evolving spatial amplitudes, phases, and Poynting vectors). This diffractive propagation property has been correctly captured by classical diffraction integrals, which form the very foundation of classical optics since its inception in 1817 by Fresnel [6].
In a typical two-beam interferometer, like Mach-Zehnder, the classical expression for the superposition of two replicated output plane waves, crossing through each other at a small angle, can be expressed as the traditional Mathematical Supposition Principle:
In Figure 1, we have assumed that relative propagation delay along the two independent paths in the interferometer is zero by virtue of physical alignment when they intersect at the X = 0, the origin where the detector array is placed to register the fringes. The intensity variation is along the X-axis following the relative path delay, .
Eq. (1) represents the well-known mathematical superposition principle (SP). SP is not directly observable. Generation of observable superposition effect (SE), or data, involves a series of complex physical interaction processes. The data are generated as some physical transformation triggered by energy exchange between some interactants in our chosen instrument. The interactants are photoresponsive material dipoles and EM waves. First, the material dipoles in the detector array experience dipolar stimulations by both the superposed EM waves. The incident optical frequency ν has to be resonant with the allowed quantum transition. This dipolar stimulation parameter χ(ν) is the linear (first order) polarizability. The traditional photo effects happen with the release of quantum mechanically bound photoelectrons to free space or internally to the conduction band. The older method used to be the dissociation of quantum mechanically bound Ag-Halide molecules in photographic plates. Therefore, the physical amplitude-amplitude stimulation effect, , should be explicitly recognized, as it is done by various semi-classical models [7, 8, 9]. Therefore, the real physical superposition principle should be expressed as the joint dipolar stimulations:
(We are assuming here that the incident light beam is linearly polarized. Further, we are neglecting interactions of light with any anisotropic medium. This is to avoid unnecessary complexity in mathematics, which can divert our attention from the simple core issue that superposition effects become manifest only when some material medium undergoes observable physical change under the simultaneous joint stimulations by multiple waves.)
The interference fringe data are generated after the energy transfer process is executed by the stimulated photosensitive dipoles, which is the square modulus of Eq. (2):
This nonlinear square modulus interaction process cannot be executed by superposed linear fields in the absence of interacting materials. Accordingly, Eq. (1) does not represent any physical interaction process. Therefore, we should restrain ourselves in assigning interpretations about the physical nature of light using this unobservable amplitude, Eq. (1), while, holding in mind, the observable data represented by the energy Eq. (3). The proper linear physical superposition principle, representing joint amplitude stimulations of a detector, should always be represented by Eq. (2) to avoid making noncausal interpretations of observable superposition phenomena of nature. The best rational and physical interpretation that we can assign to Eq. (1) is that electromagnetic waves can co-exist within the same physical volume, while they cross-propagate, or co-propagate, independent of each other, as long as the medium is linear and noninteracting. This is the generalized noninteraction of waves (NIW), mentioned earlier valid for all propagating wave phenomena. For details, see Refs. [1, 10].
Let us appreciate that the same single photon cannot represent two different physical beams. Let us also note that in Eq. (3), the fringe visibility, , is influenced by the two different amplitude parameters, and further, the fringe oscillation is influenced by two different phase factors, and , which are manifest in the oscillation of the fringes through . We have mentioned that no stable elementary particle can carry more than one unique value for any of its physically essential (critical) dynamic parameters at any moment that define the existence of the entity. Thus, as per our causal equation, each one of the two signals we generate in an interferometer must be a physically real and independent physical signal, whether they are interpreted as classical EM wave packets or quantum mechanical "indivisible light quanta." Each wave plays separate important role in generating the final fringe pattern. Just one "photon," from one or the other wave, cannot dictate the emergence of the fringes as it does not have the other necessary parameter to interact with while stimulating the detecting dipoles. The concept of interaction-free superposition effect does not advance our intention of deeper enquiry of phenomena in nature. Natural phenomena happen only through diverse interactions between different entities.
Fringes of perfect unit visibility are almost impossible to achieve in practice. The fringe visibility is , only when accurate to a tiny fraction of the energy carried by a single photon, (for green light). This is a staggeringly small energy. We do not have any energy meter that can even come close to directly measure such a miniscule energy decisively. Further, the final beam combiner must have exactly T/R = (0.5/0.5), accurate to the single photon energy. A vendor will never agree to sell such a beam splitter, as they do not have the necessary technology to measure energy with such an accuracy. In other words, the fringe visibility, , must be computed by separately measuring the two intensities, and , for the two real beams and then compute a1 and a2. Eq. (3) has been experimentally validated innumerable times during the past two centuries. This experimentally valid mathematical Eq. (3) dictates that the superposition of two real and separate beams generates the interference fringes, and they cannot be generated by a single photon out of only one of the two beams.
Emergence of dark fringes: To strengthen the previous points further, we explore the physical processes behind the emergence of alternate dark fringes between the bright fringes. Mathematically, perfect dark fringes imply . Superposition effect is local since the bright and dark fringes continuously vary in space and the size of the detecting dipoles is miniscule (∼Angstrom3) [11]. We all accept that the cosine intensity variation in the fringes is due to the phase factor , as we keep changing the relative path delay . is a minimum whenever becomes an odd multiplier to , making . However, the only way could be exactly zero is when , making the visibility factor . It is then obvious through the rigorous mathematical logic of Eq. (3) that dark fringes cannot be due to "nonarrival of indivisible photons" [12, 13] in these locations. Implication of "single photon interference" is that the detector's role is only to register the arrived energy without any participating in any way in the superposition interaction process itself. This noncausal assumption has led to the emergence of the belief behind nonlocality, noncausality, etc. This faithful belief in nonarrival of indivisible photons or of particles at dark fringe locations still dominates our physics community. It is then a fair question to ask whether we should trust more than a century old data validated Eq. (3) or the unnecessary mystical interpretation proposed after the advent of quantum mechanics in 1925. In the face of the reality that all theories are approximate and incomplete, we can keep on advancing science only by accepting any experimentally validated mathematical theories, which do not promote noncausal and incongruent interpretations. We should use such "evidence-based theories" as intermediate steps toward constructing better and better theories. In other words, QM is incomplete since it has been constructed to predict only the final measurable outcome without any attention to understand the detailed interaction processes that precede the QM transitions.
Most of the self-contradictory interpretations of the superposition principle will become unnecessary once we accept the hybrid model for photons. Atoms and molecules do emit discrete quantity of energy during each allowed quantum transition. However, the packet of energy immediately evolves into a perpetually propagating EM wave while spreading out diffractively, as it has been modeled classically (Ch. 10 in Refs. [1, 14]). During energy absorptions, the quantum dipoles function as "quantum cups" and they fill up that cup with the required amount of energy out of the classical waves propagating through them. If the flux density within its stimulated quantum-cup volume is below , then there will be no photoelectric transition. This is a difficult experiment since one has to reduce the energy flow in reliable calibrated steps toward the single photon energy of . Current technology does not offer us any energy meter that can directly measure such a miniscule amount of energy. The release of a single bound photoelectron does validate the absorption of amount of energy, since QM is essentially correct. However, it does not guarantee the existence and presence of indivisible photons. Each measured click out of our photon counting electronic system represents a burst of current pulse consisting of hundreds of millions of amplified electrons. As electrons are quantized particles and always bound quantum mechanically, their emission process steps will have to be discrete. Discreteness of the emitted electrons does not automatically validate the discreteness of the propagating EM wave energy. Quantum formalism does not require that quantum transition can be affected only by another quantum entity having the right amount of energy to donate. Striking stones create sparks. This is a complex energy exchange process, but nonetheless, it consists of a discrete series of quantum excitation and de-excitation processes triggered originally by the mechanical energy of hands. The validity of Boltzmann's rule of population density, dictated by classical thermal collisions, is rigorously obeyed by quantum systems that experience thermal collisions.
The author believes that nature abhors magic and mysticism. Physics has been advancing based on hard causality guided by rigorous mathematical logics but as a guiding tool only. Human invented mathematics cannot dictate nature how she ought to behave.
We have already defined real physical superposition principle as Eq. (2), which represents the linear amplitude stimulation of the detecting dipole induced by both the superposed fields simultaneously. The energy transfer to the detector is given by the square modulus of Eq. (2), as presented in Eq. (3). Note that the polarizability χ(ν) in Eq. (2) and χ2(ν) in Eq. (3) can be taken out of the amplitude and the intensity terms, respectively, as a common factor, provided χ does not vary with ν and t. This is allowed by the generic mathematical rule. This mathematical rule, while bears out in many situations, deprives us, appreciating the deeper interaction processes in nature. The t-independence of χ(ν) is almost guaranteed whenever the material properties are stable in time. However, the frequency independence is never rigorously valid, since response of all materials depends upon the resonant frequency (quantum transition levels or bands in atoms and materials). We know that the detectivity of broadband solid-state detectors varies with frequency. However, when the spread of the wave frequency in the incident beam is extremely narrow, χ(ν) can be treated as a constant for that frequency and treated as a constant common factor.
Let us now pay close attention to the third line of Eq. (2). With χ(ν) taken out as a common factor, the expression within the parenthesis may imply that the two EM wave amplitude components are summing themselves to create the interference of waves. We can easily make this very serious conceptual mistake guided by the allowed abstract mathematical rules. Mathematical rules are human-invented logics. They do not always capture nature's logic. The linear superposition of amplitudes, as already underscored, does not generate any observable effect by themselves, in the absence of some interacting materials. This is why appreciating noninteraction of waves (NIW) is of such critical importance [1].
The interferometer arrangement we have utilized in the previous section (Figure 1) could be described as an interferometer in spatial-fringe mode [15, 16]. The Poynting vectors of the two propagating beams in Figure 1 are noncollinear. In this section, we analyze an alternate arrangement where the Poynting vectors of the two pairs of output beams, generated by the output beam combiner BC, are perfectly collinear and coincident to each other. Such superposed parallel beams will continue to propagate with the same relative, but fixed, path delay generated within the interferometer (Figure 2). To keep the mathematical analysis simple, as in the last section, we are assuming that the relative physical path delay between the two beams is zero on arrival on the proper dielectric-coated surface of the BC. To observe energy oscillating fringes under this condition, one has to scan one of the two mirrors with a piezo-electric drive to introduce the temporally oscillating path delay . Such interferometer arrangement, in scanning-fringe-mode, is quite common in today's lab.
As argued in the above section, the observable superposition effect can become manifest only through light-matter interaction. For a scanning-fringe interferometer, the interaction is between the dielectric boundary and the two superposed wave amplitudes from the two opposite sides of this boundary material layer. This is a pure classical boundary value problem for EM waves, which was actually solved by Fresnel even before the development of Maxwell's wave equations. Fresnel found that for the external reflection (towards the direction of lower refractive index), the wave amplitude suffers a phase shift [1, 17]. The light-matter amplitude-amplitude stimulation is facilitated by the classical linear bulk polarizability of the dielectric boundary materials. Here, this is for bulk classical dipole polarizability. It is different from that for dipoles undergoing QM transition, as in Eqs. (2) and (3), even though they are both linear. Then, the joint stimulation of the boundary layer induced by the right-going and the up-going beams, respectively, is:
Assuming that the dielectric boundary layer is approximately loss-less, we can assume that . Then, the detectable energies in the right-going and up-going beams, respectively, are:
From the very last lines of the energy, Eqs. (6) and (7), one can easily recognize that, under the ideal conditions of , the sum total energy of the two beams, incident on the beam combiner from the opposite sides, remains constant, even though each output is oscillating between zero and :
The implication is that a passive 50% beam splitter can dynamically keep changing its effective physical transmitivity, or reflectivity, between zero and one. The photo at the center of Figure 2 is an experimental demonstration of this fact. A dual trace oscilloscope simultaneously displays the output of D1 and D2. While they are undergoing sinusoidal oscillation in displayed energy, the two traces are complementary to each other, preserving the rule of energy conservation. This simple classical superposition effect does not have any connection with quantum physics, since the beam splitter is purely a linear classical optical element. It is not undergoing any quantum transition or generating the fringes due to quantum absorption of energy. As mentioned before, the readers should note that currently we do not have technology to precisely set the conditions and accurate to that of a single photon energy, . Accordingly, the claims that one can let a perfectly measured single photon incident on a beam combiner and then detect its appearance in either the port D1, or the port D2, must be taken with a "grain of salt." Recall that our counting ability of quantum mechanically released electrons does not make EM radiation quantized. Dirac's quantization of EM wave cannot localize photons; however, we have been generating highly localized, in time and space, laser energy as femto second pulses. The photons as linear superposition of finite or infinite set of Fourier modes are not a causal model of physics because of noninteraction of waves.
The conservation of energy remains true even when the conditions and are not met. From the sum of the second lines of Eqs. (6) and (7), one can again derive that the total energy is still conserved:
From the standpoint of classical physics, the π phase shift for all external reflection is of critical significance behind the capability of a beam combiner to re-direct energy from one beam to the other. In fact, a homogeneous boundary layer always behaves like an "active anisotropic" layer because the boundary layer molecules are constrained to oscillate more easily as dipoles along the plane of the boundary, than orthogonal to the boundary, when an incident oscillating electric vector stimulates them. This is at the root of Brewster's Law, Malus's law, and partial polarization of "unpolarized" light in reflection. EM waves always stimulate the material dipoles to oscillate along the direction of its E-vector, while the wave propagates through any medium [17].
The π-phase shift in the external "reflection" (see the sketch for the BC in Figure 2, thick line from the bottom on the BC) means that the surface dipoles are oscillating in such a way as to allow only π-shifted light to be reflected compared to the incident beam. If a collinear "transmission" beam (Figure 2, thin line from the left on the BC) wants to pass through in the direction of the external reflection (to the right), it has to match this π-phase shift. However, if the phase of this left coming "transmission" beam happens to be zero or modulo-2π, its attempted dipole oscillation will be opposed by the "reflection" beam. The two incident amplitudes are now competing with each other in opposite directions. If the two amplitudes are equal, but opposite in phase, then all the energy of the two beams will be redirected along the upward direction, as this direction provides an in-phase path for both the beams and vice versa for the other direction.
The key point is that all beam combiners in all two-beam interferometers function this way when the Poynting vectors for the output beams are set for perfect collinearity and spatial coincidence. The simultaneous physical presence of both the interfering beams from the opposite sides is must for the superposition effect to become manifest. In other words, even if light consisted of "indivisible single photons," we must have the simultaneous presence of two photons from the opposite sides of the beam combiner. Hence, if "indivisible photons" were the reality, then such a classical interference effect would not have been observable with the incidence of truly "one photon at a time." The experiment we have presented used a 5-mW He-Ne beam, not "single photons." Such an experiment is being carried out in many senior level physics/optics laboratories.
The following challenge unequal-beam-energy experiment which is yet to be carried out. The superposition conditions can be dramatized even further. Even if one uses a beam combiner with (the energy must be conservation rule is preserved, ), one can then alter the incident energies and proportionately to completely suppress the energy propagation in one of the two chosen direction under the relative π-phase shift condition for that particular direction. Would one be able to explain such an unequal-beam-energy experiment using the model of "indivisible single photon interference"? Remember, the mathematical model consisting of the equations given above corroborates the measured data. Note that this generic mathematical model requires the simultaneous presence of the two signals from the two opposite sides on the beam combiner incident with the right phases and right amplitudes. Then only they can create the desirable transmission (or reflection) along either of the two output ports.
So far, we have only made passing negative comments about Einstein's 1905 of "indivisible light quanta" without presenting a better model to understand the interaction processes behind the photoelectron emission. In this section, we "derive" the photoelectric energy equation using the inevitable superposition of amplitudes of multiple wave groups. As before, we are following semi-classical model for light-matter interactions.
Classical thermal radiation consists of random wave groups emitted spontaneously with random phases. They keep propagating while diverging out due to waves' intrinsic diffraction property. The total dipolar amplitude stimulation of a bound electron can be expressed as:
The electron binding system must absorb the necessary quantum-cup-filling energy, before the electron can be released to the conduction band, or become a free-space electron. This is a quadratic process:
We need to collect an ensemble of data to validate any characteristic behavior of a physical system. A single event (data point) is not sufficient to verify a theory.
Note that we have "recovered" Einstein's photoelectric energy equation out of dipole amplitude stimulations due to multitudes of waves. The left curve in Figure 3 was already published for Einstein to read [18]. His brilliance was to correlate the "minimum frequency" below which no photoelectron emitted to a unique "quantumness" in the phenomenon. He assigned this "quantumness" to "indivisible light quanta" in 1905 ("photon" word was coined in 1925 by G. N. Lewis). This was a brilliant idea at that time because even Bohr atom model with quantum mechanically bound electron was published 8 years later in 1913. However, had Einstein correctly assigned the quantumness to bound electrons, he would have formulated mathematics of quantum mechanics in his own style some 20 years earlier. Unfortunately, this neglect of the physical importance of amplitude stimulations as the critical first physical step before any quantum transition (energy exchange) can take place continued even after the QM was developed in 1925 and formalized during the next 10 years. We are stuck with the fictitious "indivisible light quanta" and "single photon interference." It would be a great cultural anthropological study to analyze and understand why the absolute majority bend over backward to justify Einstein's mistaken postulate, even though we now know that all electrons are bound quantum mechanically in materials. Recall the third line of Eq. (3). Waves only fill up the quantum cups with the necessary energy if the dipoles are resonant to the frequency of the incident waves (see left-diagram in Figure 3).
Let us rewrite Eq. (12) under the assumption that incident light is coming from a single frequency laser. In that case, can be considered to be a constant and can be taken out of the summation sign:
The last step in Eq. (13) is meant to underscore again how some human invented mathematical logic, while corroborate measured data numbers can misguide us to believe that waves by themselves can carry out the complex operations of first (i) summing the wave amplitudes, and then (ii) taking square modulus of the sum.
In both Sections 2 and 3, we have underscored the significance of noninteraction of waves (NIW). The neglect of NIW made us culturally oblivious to the fact that the linear mathematical superposition principle (SP) is not an observable phenomenon of nature. It does not embody any interaction exchange process followed by nature. In contrast, the superposition effect (SE) is an observable phenomenon because it models light-matter interaction process through the nonlinear quadratic operation . Note further that SE becomes manifest only in material detectors when their frequency resonant property matches with the frequency of the incident wave. NIW was known at least about 1000 years back [2]. It was repeatedly re-discovered but was neglected [10]. The puzzling issue is that this NIW is mathematically built into Huygens-Fresnel diffraction integral. Many modern undergraduate texts show constructions of Huygens' secondary wavelets, evolving through each other unperturbed, to model diffraction, reflection, and refraction, while propagating through linear and anisotropic media [17]. Yet, current books and literature never mention NIW. However, they present the linear superposition principle as the summation of wave amplitudes as if they physically sum themselves even while explaining that all propagation of light involves stimulating the dipoles of the material media. Note that the physical summation of wave amplitudes of different frequencies is behind the derivation of "group velocity," which is not an allowed physical process of nature. Because of the flexibility in interpreting math, our time-integrated data gathering practice, etc., many of the data validate the mathematical model. This is how we have learned to accept SP as an observable phenomenon by itself through years' of training. In the process, we have learned to suppress our inherent enquiring minds regarding the obvious contradictions while explaining many optical phenomena (see Ch. 1 in Ref. [1]). How have we acquired this culture of deferring to math and data without enquiring about the physical interaction processes? It took a series of grand mathematical successes in physics, during late 1800s and early 1900s, provided by only math and data, which is now known as the "evidence-based science." Evidence-based (experimental) science is the best possible approach to generate objective information about natural phenomena. In addition, mathematics is the best possible logical and objective tool, so far invented my humans, to model nature. Unfortunately, neither of them separately, or jointly, can provide us with the complete information about any interaction in nature that we study. We have learned to ignore the fact that, albeit math being the best objective logical language to model nature's behavior; it is just another human-invented language. Further, causal (or objective) mathematical equations require interpretations by human subjective minds. Math does not represent logical language that nature is obligated to follow. We have also learned to over-ride the fact that evidences (experimental data) still suffer from: (i) competing interpretations of data assigned by different subjective human neural networks, which have actually evolved for biological survival and not for the objective determination of the rules behind the biospheric and the cosmo-spheric evolutions. (ii) Further, no finite set of experimental data can provide us with the complete information about any natural entities. To be objective, we need to put careful and conscious efforts. Since the very beginning of biological evolution, our pragmatic system engineering approach to solve problems at hand with limited knowledge and trial-and-error has already made us the "top-dog" in the biosphere. We have installed the Global Internet System by figuring out the core system engineering functions for (i) generating, (ii) modulating, (iii) propagating, and (iv) detecting electrons and photons. However, we still do not understand what "photons" and "electrons" are made of. The lesson-to-be-learned is that a system engineering approach keeps us both grounded and pragmatically advancing. Whatever tools and technology works, they really obey the laws of nature even if we cannot articulate the rules.
Unfortunately, over the last century, we have deviated from consistently applying the system engineering approach to understand fundamental processes in nature [19]. This is evident from the broad cultural acceptance by the Knowledge Gatekeepers that the theories of Relativity and the mathematical formalism of Quantum Mechanics provide us with the final foundation of all future physics. Fortunately, the very long-term history of science does recognize that all human constructed theories are necessarily incomplete since they have been formulated based upon insufficient knowledge of the universe.
During the formative years of QM, people were marveled by the fact that Schrodinger's always gives the right result even though they could not give proper physical explanation regarding what physically represents. Born explained it as the abstract mathematical "probability amplitude," which must undergo an instantaneous "wave function collapse" to generate data in our instrument. This is certainly not a causal approach to understand and visualize the real physical interaction processes in nature, which gives rise to observable physical transformations. This conceptual gap between and gave birth to the concept of "measurement problem," which was then "solved" by using various elegant mathematical theorems. However, we have ignored the fact that if gives us real world data, then must also contain physical realities embedded in it. The author believes that represents the real physical amplitude stimulation of the entity under study, as presented here in Sections 2 and 3 (see Ch. 3 in Ref. [1]). The square modulus operation represents energy exchanging interaction process between the interactants and between the energy donor and the energy recipient, guided by a force of interaction. These data generating interaction processes take a finite time. The interactants must first find the interaction compatibility with each other guided by the force they are experiencing and then undergo the full stimulation before jointly executing the square modulus operation to exchange the quantum cupful of energy. Let us elaborate the logical steps behind this physical process (see Ch. 12 in [1]).
Data are physical transformations: A properly designed instrument can register the anticipated data only when the measuring instrument receives the appropriate signal indicative of some physical transformation experienced by the chosen interactants.
Physical transformation require energy exchange: Data generating physical transformation can happen only when the interactants exchange the necessary energy.
Some force of interaction guides the energy exchange: Only an allowed force of interaction, compatible with the interactants, can guide energy exchange toward some physical transformation.
All "consumed" interactions are necessarily local: Since all forces are of finite spatial range, the interactants must be physically present within this range of the guiding force. Thus, all interactions are "local" for the interactants as they must be within each other's vicinity to experience the mutual force, which "dies out" with distance. This implicates that interacting particles cannot remain "entangled" beyond the range of the effective force of interaction (in the sense of continued mutually influencing remote interaction). Nature is strictly causal, not mystical. There is no interaction free superposition effect.
The purpose of Physics is to figure out the invisible interaction processes that
generate the data: Our evolution and survival in the biosphere demand of us to think like system engineers and learn to manage the complex biospheric system. Unfortunately, we are not teaching our younger generation the extreme importance of understanding the depth of interaction processes going on in nature.
The information retrieval problem: We can now appreciate that the validation of a theoretical model for any specific interaction by some experimentally measured data does not help us visualize the microscopic interaction processes that generated the data. This is an eternal "Information Retrieval Problem" for humans. This is not a "Measurement Problem" that can be solved by further refinement of the instrument or by inventing some elegant mathematical theorem. Nature is a creative system engineer. From this standpoint, we must figure out how to understand the details of the interaction processes. Today, we are facing the challenge of geoengineering the biosphere to slowdown the oncoming global warming. This is a formidable system-engineering task. Unfortunately, the physicists cannot contribute in this urgent process unless they are trained to enquire and understand the invisible interaction processes as system engineers.
How we can attempt to visualize the invisible interaction processes: First, we need to change our culture to accept explicitly that "evidence based science" will always represent incomplete science. Only our consistent enquiry in understanding the interaction processes will allow us to start getting closer and closer to understand nature's system engineering marvels at both the micro and the macro levels. At present, we are happy with the Measurable Data Modeling Epistemology or MDM-E. MDM-E has successfully guided us to the highly advanced state we are in today. The author is proposing that we add Interaction Process Mapping Epistemology, or IPM-E, in conjunction with the prevailing MDM-E and accelerate our deeper understanding of nature as creative system engineers. Since the deeper realms and reality of nature are not directly visible to us, we will have to keep on applying IPM-E iteratively, perhaps, indefinitely. The progress will be slow, but we certainly will not fall in blind love with theories that are validated by data, but nonetheless do not represent nature's actual working rules. Slowly, we will become efficient and conscious sustainer of the biosphere, instead of continuing to be the plunderers. Declaring war against the evolutionary behavior of nature (outcomes of the laws of nature) will be futile until we master the extremely complex system engineering rules of the biosphere. This is why we must teach the students to keep on challenging the foundational postulates behind all working theories and assure consistent evolution in scientific theories. Sections 2 and 3 of this paper represent out of measurable data for superposition effects.
Let us recall Newton's philosophical view toward the end of his life: "If I have seen further than other men, it is by standing on the shoulders of giants." Since 1400 until today, we must have been accumulating new knowledge developed by many thousands of deep thinking scientists. Conceptually, we have built a pyramid of knowledge out of all these contributions. We should learn to climb to the apex of this pyramid to increase our knowledge horizon. We should not bow down our head out of "messiah complex" and reduce our knowledge horizon.
Einstein is known to have said: "…. After 50 years' of brooding over the question of what are light quanta; I still do not understand it!" This is the sign of a truly honest enquiring mind of a scientist. In spite of receiving the Nobel Prize for his proposed concept of "indivisible light quanta," he kept on questioning it due to causal inconsistencies. This public "questioning" attitude of Einstein has provided the author the strength to explore critically whether "indivisible light quanta" can really exist [20]!
Our key objective behind this chapter has been to re-introduce noninteraction of waves (NIW), which is a very important reality of nature in the field of interferometry. Since different "photon" amplitudes cannot interact even while co-propagating or cross propagating through each other, once spatially separate, EM wave packets cannot keep on influencing each other remotely; in this causal sense, they cannot remain "entangled." I leave it to the readers to make the necessary adjustments in their future thinking whether "single photon interference" truly represents the reality of nature [21, 22].
The readers should also note that the implication of the generic NIW-property of all waves is very significant in most of the branches of physics, some of which have been elaborated in my book [1]. I believe that incorporation of NIW in our analytical thought processes would help us develop deeper understanding of nature, besides guiding us to improve upon and/or invent many new instruments.
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