Advances in Neuroimmune Biology 5 (2014) 75-89DOI 10.3233/NIB-140085IOS PressPhysiological, Immunological andEvolutionary Perspectives of Labor as an Inflammatory ProcessJosé Javier Reyes-Lagosa, Juan Carlos Echeverría-Arjonillaa, Miguel /Ángel Peña-Castilloa,Augusto J. Montiel-Castrob,c and Gustavo Pacheco-Lópezc’a Electrical Engineering Department, Universidad Autonoma Metropolitana (UAM) Campus Iztapalapa,Mexico City, Mexicob Darwin Center for Evolutionary Thinking (CEDAR), Philosophy Department, Social Sciences and HumanitiesDivisión, Universidad Autónoma Metropolitana (UAM) Campus Iztapalapa, México City, MéxicocHealth Sciences Department, Universidad Autonoma Metropolitana (UAM) Campus Lerma, Lerma, MexicoAbstract. The precise mechanisms for the onset of labor at term remain unknown, yet several studies in humans reveal the role of cytokines in the initiation and maintenance of labor, showing many of the hallmarks of inflammation. Recent findings suggest a possible relationship between the activity of the autonomic nervous system (ANS) and the vagal anti-inflammatory response during labor. Furthermore, the role of vaginal microbiota is particularly important during pregnancy because vaginal dismicrobism is one of the most important mechanisms associated with preterm birth. In this review, we present evidence suggesting that a sterile anti-inflammatory response is manifested to attenuate the excessive inflammation introduced by low-risk labor at term, involving either the action of a cholinergic pathway, uterine-like myokines or the vaginal microbiome.Keywords: Anti-inflammatory response, heart rate variability, vaginal microbiota, pregnancy, labor7029452858770ISSN 1878-948X/14/$27.50 © 2014 - IOS Press and the authors. All rights reservedThis article is published online with Open Access and distributed under the terms of the Creative Commons Attribution Non-Commercial License.ISSN 1878-948X/14/$27.50 © 2014 - IOS Press and the authors. All rights reservedThis article is published online with Open Access and distributed under the terms of the Creative Commons Attribution Non-Commercial License.INTRODUCTIONHuman pregnancy and labor are physiological scenarios during which the fetal allograft must be tol- erated, fed and allowed to grow. The process of labor is mounted to deliver the fetus from the woman's uterus [1]. Parturition is characterized by both cervical ripening and myometrial maturation followed by uterine contractions leading to cervical dilatation and birth [2]. Given that labor seems to be driven by pro-inflammatory cytokines, we suggest that it shows inflammation hallmarks and that the immune privileges that the fetal-placental unit enjoyed during pregnancy are suppressed at the time of labor [3].During term and preterm labor different immuno- logical changes are observed, including acute inflammatory features such as an increased influx of leucocytes and an elevated expression of pro- inflammatory cytokines that have been observed in cervical tissues and fetal membranes [4-6]. Likewise, many studies support the hypothesis that labor involves an inflammatory event [7-13], but more precisely, that human labor should be considered as a sterile inflammatory event mainly because it may be free of microbes triggering labor. According to Rock et al. [14], some sterile pro-inflammatory stimuli, like dead cells, irritant particles (including crystals, minerals and protein aggregates), trauma, immunogenic antigens and autoimmune conditions can cause an inflamma- tory response. An important finding was that all of such stimuli elicit this inflammation through the same cytokine mediator, interleukin-1 (IL-1) [14]. Given that labor implies substantial muscular work involving sig- nificant physical efforts, with an important metabolic activity [15, 16], we hypothesize that rigorous energy regulation must then be achieved by homoeostatic physiological processes during this final stage of preg- nancy where inflammation is also occurring.Other authors have reported that an inflamma- tory input activates fast and subconscious anti- inflammatory neural responses [17]. In fact, Huston and Tracey [18] defined a neuronal circuit acting as the anti-inflammatory reflex in which the main component is the descending branch of the vagus nerve. Centered on this model, action potentials coming from vagus nerve lead lymphocytes to release acetylcholine that inhibits the production of pro-inflammatory cytokines through a7 nicotinic receptors [18,19]. This consider- ation then leads to propose that the observation of the activity of the vagus nerve can provide a suitable tool for measuring an anti-inflammatory reflex in various physiological contexts, thereby supporting a relation- ship between the activity of the autonomic nervous system (ANS) and an anti-inflammatory response [18]. A conventional noninvasive method to assess the ANS is by analyzing heart rate fluctuations (HRF) data [20]. In general, linear and non-linear methods are applied to obtain measures that are used to estimate parasym- pathetic and sympathetic activities [20]. Additionally, some studies reinforce the consideration that the car- diovagal activity, quantified by the analysis of HRF data, could provide insights to understand the inmuno- vagal route [21].In addition, the role of vaginal microbiota in relation to the process of labor may be particularly impor- tant because vaginal dismicrobism is one of the most important mechanisms associated with preterm birth and perinatal complication [22]. Some authors have reported that a probiotic dietary supplementation can be related to a global anti-inflammatory effect on the vaginal immunity, with potential implications in preventing preterm birth [23]. However, most of the vaginal microbiota of pregnant women has been stud- ied during pathological situations such as bacterial vaginosis and aerobic vaginitis [24-28]. As the role of microbiota should not be limited to pathological conditions, we consider necessary to reevaluate the physiological role of vaginal microbiota during preg- nancy and its relationship with inflammation.Current research on the microbiome is exposing several dependencies of the host on microbiota for the maintenance of health, and also the crucial role that endosymbionts may play in the initiation and propagation of disease. Thus, several experimental model systems, including mice, fish, insects, squid and non-human primates, continue providing impor- tant insights about the host-microbiota homeostasis [29, 30].Focused in the physiological process of labor at term, the purpose of this review is to address the fol- lowing questions linked to labor in healthy women: Is an anti-inflammatory reflex an adaptive response to cope with the excessive inflammation occurring during labor? Could this process be assessed by the analy- sis of heart rate dynamics, immunological biomarkers and the characterization of vaginal microbiota? To pro- vide some insights into these questions, first we review important issues on the energetic homeostasis during labor. Then, bypresenting evidence about theincreased vagal modulation during labor at term indicated by HRF data, we focus on the sterile anti-inflammatory reflex. Finally, bymeans ofacomparativeapproach, we present a few evolutionary aspects of the relationship between vaginal microbiota and pregnancy.ENERGY HOMEOSTASIS DURING LABORDuring pregnancy, the fetus requires glucose and amino acids for its growth and development, which sets the pregnant woman to a continuous state of energy- demand. Studies of protein and energy metabolism demonstrate the need of regulating the usage of those nutrients [31, 32]. Fat is deposited in maternal stores throughout pregnancy, providing an energy reserve of 30 000 kcal at the end of gestation [31]. Of course this reserve is important to satisfy the fetal requirements, but possibly, also, to confront an important period of energy expenditure caused primarily by an increasing contractile activity in the myometrium during labor. In fact, the maternal heart rate during labor increases up to levels comparable to moderate or heavy exercise [33].The molecular mechanisms involved in the uterine quiescence during gestation and the induction of labor are not completely known, though it has been demon- strated that the energetic metabolism of the uterine muscle is different between pregnancy and labor [34]. As glucose represents the primary maternal energy reservoir during the first and second stages of labor [35], if labor is prolonged and the source of glucose is limited, gluconeogenesis can be insufficient. Dur- ing normal labor, the concentration of free fatty acids and ketone bodies increases, leading to the recruitment of substrates other than glucose and a relative reduc- tion of carbohydrates [36, 37]. Thus, labor's efforts rapidly decrease the availability of carbohydrates, lead- ing the women's body to metabolize fat for obtaining energy. Consequently, the availability of amino acids in the mother's and fetus' bloodstream is reduced, while the fatty acids and ketones increase [38]. Furthermore, Scheepers et al. suggest that maternal hyperglycemia occurs during labor, leading to increments in maternal and fetal lactate production, resulting in metabolic acidosis [35].It is well known that, during physical activity, the consumption of energy is linked to the actual dura- tion of this activity [39]. But, given that labor can be extended for several hours (the average time of labor for a primiparous woman is about 16 to 17 hours) [40], it becomes a high energy-demand process similar to that of mid-heavy exercise according to Sohnchen et al. [33]. One may ask how much energy is being used during labor. For instance, ifthe energy expenditure for a 55 kg woman during moderate and heavy exercise is between 3.5-7.7 Kcal/min [41], therefore the estímate of energy consumption is 3400-7900 Kcal for a woman during 16 to 17 hours in labor. However, there is few empirical information about energy-expenditure dur- ing labor. It has reported as low as 50-100 Kcal per hour of active labor [42] and as high as 300-520 Kcal, Maganha and Peracoli [43]. Why is this big varia- tion occurring? One likely answer may be related to the differences in energy expenditure between the first and second stages of labor [44]. However, Katz et al. have reported that due to the intermittent character of uterine contractions, normal labor and delivery do not impose high energy demands on the parturient, but that a prolonged labor may result in maternal metabolic disturbances [45]. Thus, these considerations provide opportunity for the realization of future research on this phenomenon.Other studies indicate that rather than lipids, the pregnant uterine smooth muscle uses glucose as its main nutritive metabolite, and that compared to the striated muscle, the anaerobic pathway of the glucose metabolism is more active in the myometrium [46]. Also, it has been suggested that glucose has a critical role as the principal fuel for ATP formation and both the adenylate kinase and 5'-nucleotidase reactions are involved in any event of glucose shortage [46]. Other studies suggest that glycogen is important because the myometrium content in pregnant rats was found to be increased just prior to parturition [47]. Findings dur- ing the latter halfofpregnancy in pregnant rats showed gradual increments ofinsulin binding sites that peaked on the day ofparturition [48]. Other findings suggested the effectiveness of insulin in the uterus of pregnant rats, showing that its sensitivity in the myometrium was similar to that observed in skeletal muscle [49]. Thus, myometrium inflammation by means of insulin resistance may be required for maintaining the flow of energy that sustains uterine contractions.From an immunological pointofview, labor can also be considered as an inflammatory event [7-13]. From this perspective, the regulation of energy homeostasis and the immune response is crucial for an organism's survival [50]. For example, in chronic inflammatory diseases, balanced energy-rich fuel supply is largely disturbed due to the vast consumption of an activate immune system [50]. In turn, this process must be occurring during labor as well to fulfill the energy con- sumption of the mother and fetus. Thus, the maternal immune response and the metabolic regulation should be integrated and the proper function of each must be highly dependent on the other. According to Hotamis- ligil et al. this interrelation can be viewed as a central homeostatic mechanism [51]. It is well known that an immune response involves a substantial investment of energy [52-54]; therefore, immunity is dependent on a trade-offbetween other highly demanding energy pro- cesses [55]. Labor is a process of energy consumption where the energy-immune trade-off between mother and child is still unclear. Thereby, from this point of view, labor can be regarded as an ‘energy demand reac- tion' caused by the activity of the immune system, the mobilization of fuel stocks (lipolysis/glucogenolysis), and the induction of insulin resistance in liver, adipose tissue and skeletal muscle with the overall purpose of allocating energy-rich fuels to the activated immune cells [50] and, clearly, also the myometrium.An orthodox trigger inflammation is infection, with the inciting stimulus being certain molecules of the invading microbes. However, several other stimuli, including mechanical trauma, ischemia, toxins, min- erals, crystals, chemicals, and antigens, also trigger inflammation in a so called sterile fashion [14];thereby, we presume that a possible phenomenon of sterile inflammation could be labor itself. Considering that most of the signs and symptoms of inflammation are caused by changes in the local vasculature of an affected tissue [56], in case of spontaneous deliv- ery and preterm labor, there are changes mediated by inflammation in the cervix tissue and the myometrium membranes [3, 4]. As the immune system needs many energy-rich supplies [57], local inflammation must be supported by fuel provision from local or systemic stores. Thus, the participation of an anti-inflammatory response is vital to restrain inflammation [17] and to limit the energy demands.This situation then raises the question: Is it possible to identify a sterile inflammatory response during low- risk labor at term in women?Any answer to this question should first consider that labor becomes exacerbated by inflammatory process. In fact, pro-inflammatory cytokines such as interleukin: IL-1, IL -6, Tumor Necrosis Factor (TNF-a), IL-8, IL- 16, Monocyte Chemoattractant Protein 1 (MCP -1), Macrophage Inflammatory Pro- tein (MIP- 1a), and chemokine CCL5 are all known to be involved as responsible factors for preterm labor [58]. Furthermore, these cytokines are present in high concentrations in the amniotic fluid of patients with infection or inflammation, stimulating uterine contractility by inducing prostaglandins. As it has been demonstrated that the activation of Nuclear factor Kp (NF- Kp) in human myometrium up-regulates inhibitory progesterone isoforms [59], some studies have found that the NF- Kp is a key modulator of these pathways and acts by regulating the expression of prostaglandins, chemokines and the pro-inflammatory cytokines involved in both term and preterm labor [1, 60]. The removal of the immunosuppressive and quiescent influences of progesterone by its functional withdrawal may in fact itselfbe initiated by the inflam- mation and activation of NF- Kp [1, 61]. Therefore, they induce premature labor and birth; yet, apparently, the anti-inflammatory cytokine IL -10 has an impor- tant role in regulating the immune response because an increased concentration of it during preterm and labor at term in amniotic fluid has also been identified [58].A STERILE ANTI-INFLAMMATORY REFLEX DURING LABORAs described above, pregnancy itself involves an inflammatory process associated with increased pro- inflammatory cytokines [1, 4, 58, 60]. In this sense, it is also known that progesterone has an immunosup- pressive role which promotes the immune privileges of the fetus and regulates the levels of cytokines in the maternal-fetal interface. However, when labor is triggered, these privileges are no longer preserved, causing a sterile inflammation (dilation of cervix, effacement and uterine contractions) and producing pain [3]. Therefore, to oppose this inflammation, the anti-inflammatory cholinergic reflex, among other mechanism explained, could in principle restrain, as in other contexts, the release of pro-inflammatory cytokines [17]. In some conditions, a synergistic coactivation of sympathetic/parasympathetic systems [17] is also manifested through the release of adrenaline/noradrenaline and acetylcholine, respec- tively. Because pain and fight reflexes are probably manifested during labor, a sympathetic participation may also be involved. Thus, both systems may proba- bly act to down-regulate inflammation (Fig. 1).Considering the role of myokines [62], it is now recognized that skeletal muscles express and release myokines into the bloodstream in response to mus- cle contraction by physical activity. Given that muscle fibers express the myokine IL-6, some studies sup- port the relationship between an anti-inflammatory effect of exercise and IL-6 [63, 64] because these myokines stimulate the production of the classical anti-inflammatory cytokines IL-1ra. A direct parallel between the three major myometrial phenotypes (i.e. proliferative, synthetic, and contractile/labor) and the three phases ofimmunological transformation (i.e. ini- tiation, tolerance and activation) [65] has even been reported, thus pointing to the question over what is the participation of myometrial myokines during labor? Some reports suggest that the stretch of uterine myocytes increases the IL-8 mRNA expression during the onset of labor [66]. Moreover, that the cytokines IL-1p, IL-6 and IL-8 levels are higher in maternal serum during the onset labor than those of non-labor women [67]. By making an analogy between exer- cise and labor, we hypothetisize that the strong uterine activity induces a sterile anti-inflammatory reflex by means of myokines production.One of the major triggers of inflammation is infection, being certain molecules of the invading microorganisms the inciting stimuli [68-70]. In this context, because labor must be free of microbes that trigger it, and in comparison with some preterm labors triggered by bacterial infection [22], low-risk labor could be considered as a sterile inflammatory event. However, while the final inflammatory manifestation of labor should be similar between infectious and ster- ile triggers, the specific control of the response may be different. Thereby, each inflammatory input may activate a distinct anti-inflammatory response; in case of low-risk labor at term a sterile anti-inflammatory response. Results showed that the IL-1 pathway plays a key role in the neutrophilic inflammation to diverse sterile stimuli, including a variety of irritant particles and dead cells [14]. Interestingly, this same cytokine has been implicated in the mechanism of human par- turition, because some results showed that pregnant women without labor had undetectable IL-1 a concen- tration in comparison to a labor group [71].Fig. 1. Diagram of the inflammatory process as a mechanism present during labor and its possible association with a regional anti-inflammatory sterile response. According to Tracey KJ, 2002, this response sometimes presents a synergistic action by the sym- pathetic/parasympathetic systems to contend with inflammation. Uterine-like myokines and the vaginal microbiome may also be involved (see text). (Image adapted from Peltier MR, Immunology of term and preterm labor, 2003).INCREASED VAGAL MODULATIONDURING LABOR AT TERMA noninvasive method to assess the autonomic ner- vous system (ANS) is the analysis of HRF data. HRFanalysis aims to separate and quantify the autonomous cardiac response in sympathetic (adrenergic) and parasympathetic (cholinergic-vagal) influences. The sympathetic and parasympathetic branches of ANS and their influences on heart rate (HR) and HRF are well understood. Sympathetic activity tends to increase HR and decrease HRF, whereas parasympathetic tends to decrease HR and increase HRF [20].The time intervals between consecutive heart beats are measured in the electrocardiogram (ECG) from the beginning of a QRS complex to the beginning of the next QRS complex. They are conventionally named RR intervals (Fig. 2a). Thus, the HRF is defined as the variation of consecutive RR intervals. Among different mechanisms, this variation depends on the autonomic control of the heart.A group of researchers began a series of discus- sions about the role of HRF analysis as an index of autonomic control, concluding that different anal- ysis techniques of HRF provided indices associated with the autonomic control of the heart [72]. Recent studies have specifically linked the cholinergic anti- inflammatory activity with changes in some of these HRF indices; thereby, considered as important param- eters useful for recording the activity of such pathway [18]. In particular, given that the root mean square of the successive differences (RMSSD) is commonly used as an index of a vagal nerve mediated cardiac control, which is closely linked to respiration via the respiratory sinus arrhythmia (RSA) [73], this is one of the HRF time-domain tools that may be likely linked to the anti-inflammatory cholinergic reflex [74]. Parasympathetic activity can also be quantified by the power-spectral analysis of the HRF, providing the high frequency (HF) index. Which physiological inflam- matory processes modify the RMSSD index? For instance, some atopic diseases, such as dermatitis, have been linked to changes in the autonomic modulation towards a higher parasympathetic influence. Patients with dermatitis had higher values of parasympathetic parameters (increased RMSSD) than controls. There- fore, authors have concluded that regional adjustments should be caused by anti-inflammatory cholinergic reflex relieving dermatitis symptoms [75]. Further- more, other studies reinforce the suggestion that cardiovagal activity, quantified by the analysis of HRF, could provide insights to understand the immunovagal route [21]. Importantly, some clinical studies have also evaluated the relationship between inflammation and HRF parameters in different contexts [76-84].In arecentstudy,we explored vagal modulation dur- ing labor at term assuming that an anti-inflammatoryFig. 2. a) An RR interval is the elapsed time between two successive QRS complexes of the ECG. RR intervals show the variation between consecutive heartbeats. Heart rate fluctuations (HRF) measurements analyze how these RR intervals change over time. b) Typical data of heart beat fluctuations during the last trimester of gestation prior to labor TRD (Left) and during labor at term LAB (Right) segments. Values of the parameters HP and RMSSD calculated for those series are also depicted. Bottom of the figure illustrates how these parameters are obtained.10325106458585RMSSDRMSSD17856206546850n1n133947106458585(1)(1)cholinergic response was manifested during low-risk labor. Thus, once obtained informed consent, we col- lected at Maternal and Child Research Center, Mexico City, Mexico (CIMIGen), 10 minutes segments of elec- trocardiogram (ECG) from 30 women in low-risk laborat the semi-Fowler position (mean age of 26 ± 5 years) without any complication and with normal pregnancy outcome: mean APGAR (5 minutes) 8.9 ± 0.7 point, gestational age of 40 ± 1 weeks and birth weight of 3175 ± 297 g.Women received intravenous oxytocin during early labor to improve contractility. The ECG were recorded during periods with low uterine activity or not present- ing contractions at all (LAB group), with the purpose that hemodynamic changes due to uterine contractions did not affect the HRF measurements. A second group included HRF data segments collected during the last trimester of gestation prior to labor from a different group of 30 pregnant women (TRD group) with gesta-tional age of 35 ± 2 weeks.A ECG portable device, Monica AN24, was used for data acquisition. Sampling frequency was 900 Hz. Required segments were delineated visually by using the device's software, displaying values of maternal and fetal heart rate among ECG-derived uterine activity.Raw maternal ECGs were then processed using pre- vious validated algorithms to generate RR intervals or HRF series corresponding to LAB and TRD segments [85]. All series consisted of 600 samples (spanning 5 to 10 minutes duration).Segments of both groups were analyzed to esti- mate the average heart period (HP), RMSSD, and HF peak index to estimate the respiration rate [86]. Previous research [87] has shown that the central fre- quency location of the HF peak of HRF (Fig. 2b) can be an appropriate index of respiration rate. There- fore, the HF component of HRF is a useful proxy for respiration rate when respiration is not directly measured.RMSSD was estimated as:n-1(NNi+1 + NNi)2i=1where NNi is the duration of the i-th NN interval in the analyzed ECG, and n is the number of all NN intervals [20, 88].We found RMSSD statistical differences (p < 0.03) between LAB and TRD groups (36 ± 14 ms vs. 25 ± 13 ms, respectively), without affecting neither HP (705 ± 100 ms vs. 705± 90 ms, respectively), nor theTable 1Mean values (±SD) of parameters of HRF (HP, RMSSD and HF Peak). Gestational age of LAB (40 ± 1 weeks) vs. gestational age ofTRD (35±2 weeks) were statically different (p < 0.00001) between LAB and TRDGROUPHP (ms)RMSSD (ms)HF PEAK (Hz)LAB705 ±10036 ± 14*0.29± 0.06TRD705 ±9025± 230.29± 0.06*p < 0.03 between LAB and TRD.HF peak (0.29 ±0.06 Hz vs. 0.29± 0.06 Hz, respec- tively); (Table 1).These results indicate that the parasympathetic mod- ulation of HRF was increased in the labor group (we found higher levels of RMSSD in LAB vs. TRD groups). This parameter then indicates an increased parasympathetic activity during labor that may reflect a counter-regulatory mechanism to restrain inflamma- tion. Noteworthy, this increment did not affect the HP parameter and thus we can discard differences in the influence of ventilation between TRD and LAB groups as indicated by the HF Peak.PERINATAL MICROBIOTAIt is now known that vaginal bacteria commensals, including Lactobacillus crispatus, L. jensenii, and L. rhamnosus, have strong associations with the vaginal epithelial cells. Vaginal bacterial commensals do not trigger cytokine secretion, but as Lactobacilli play and active role to modulate cytokine production [89], bacteria non-commensals can produce an inflammatory process. Also, this consideration is supported by the fact that recent discoveries of variation in the compo-sition of the microbiome of healthy individuáis [90-92] show the importance of identifying the processes that could possibly give rise to such variation [93], and there is a rising interest to study microbial community ecology and the healthy microbiota [94, 95].Before considering pregnancy, it is necessary to review how is vaginal microbiota characterized in non-pregnant women. Studies have described the composition of the vaginal microbiota during non- pregnancy: Ravel et al. reported the results of a pyrosequencing analysis of vaginal specimens from a sample of 400 non-pregnant healthy women; they showed that the vaginal microbiota was dominated by Lactobacilli, including L crispatus, L iners, L jensenii and L. gasserii [91]. In accordance with these results, other studies support the observation that species of Lactobacillus dominate vaginal microbiota in most normal and healthy women [96-99]. However, the dynamics of the community composition of vaginal microbiota are also affected by the phase of the menstrual cycle, ethnicity, and to a certain extent, sexual activity [100]. Moreover, it must be stressed that the concept of a “normal and healthy” vaginal microbiota is hard to describe without having a full understand- ing of its role and its effect on women's physiology. According to some studies, one possible way to sepa- rate between healthy and unhealthy vaginal microbiota is based on the resulting predisposition to acquire sex- ually transmitted infections [98, 101].On the other hand, further evidence indicates that the structure of the vaginal microbiota differs signifi- cantly during pregnancy, while other evidencesuggests that the microbiota of pregnant women is less diverse and rich (in measured variance) throughout gesta- tion and in proximity to the uterus. According to Aagaard et al. the richness of vaginal microbiota diver- sity is reduced in pregnancy, with dominance of the Lactobacillus species (L. iners crispatus, jensenii and johnsonii, and the orders Lactobacillales, Clostridi- ales, Bacteroidales, and Actinomycetales [102]. In this study authors assume that the increased Lactobacillus dominance in the vaginal microbiota during pregnancy may be important for the first inoculum that determines neonat.e's upper gastrointestinal microbiota upon delivery, or for reducing the risk of ascending infection or preterm birth. In fact, these results may be linked with a remodeling of the gut microbiota over the course of pregnancy. The gut microbiota regis- tered during the first trimester is comparable to that of normal healthy controls, but this similarity shifts sub- stantially (in terms of phylogenetic composition and structure) over the course of pregnancy. By the third trimester an enrichment of Proteobacteria and Acti- nobacteria is observed in the majority of women [103]. Ottman et al. have reported that the activity and com- position of the gut microbiota is affected by genetic background, age, diet, and health status of the host [104].The possible interactions between gut and vaginal microbiota are caused by the close proximity of the vagina to the anus, exposing the vagina, cervix, and uterus to bacteria and other microorganisms, andsome- howgiving the intestine flora access to the reproductive tract via the rectum and perineum.What factors lead to this unbalance or dismicrobism of vaginal microbiota? The incidence of vulvovagi- nal candidiasis, urinary tract infection, and bacterial vaginosis (BV), among other infections, affect vaginal microbiota [105-108]. For example, during BV the microbiota is dominated by Gardnerella vaginalis [109] and a number of anaerobic organisms. In con- trast, normal flora is dominated by various Lactobacilli producing an acid environment with pH in the range of 3.5 to 4.5 [110]. Bacterial vaginosis and aerobic vaginitis have been claimed as important mechanisms responsible for preterm birth and perinatal compli- cations [22, 111]. In addition, BV flora predisposes women to infection by the human immunodeficiency virus and sexually transmitted diseases [98]. Addition- ally, some authors even suggestthat, given thehygienic practices of contemporary societies, an excessive use of feminine-hygiene products could cause imbalances in the vaginal microbiota which may promote infection [112].It is also interesting to understand the mechanisms of defense used in the control of infections; vaginal defenses include: (i) mechanical barriers such as the mucus layer, hemocidin and fibronectin, (ii) host protective enzymes (e. g. lysozyme and lacto- ferrin), and (iii) host immune responses including the complement system and secretory immunoglob- ulin A (IgA), all of which are influenced by the vaginal microbiota [113-115]. Another way to sus- tain a healthy vaginal microbiota could be based on the oral and vaginal administration of probiotics. The application of multi-strain probiotics to improve pregnancy outcomes has already been tested in a pilot study, with some success in terms of modulat- ing immune parameters and vaginal microbiota [23]. Such probiotic supplementation can be associated with a global anti-inflammatory effect on vaginal immu- nity, with potential implications for preventing preterm birth [23]. Particularly, L. rhamnosus GR-1 is able to up-regulate the anti-inflammatory cytokine inter- leukin 10 (IL-10) and the colony-stimulating factor 3 (CSF3); and, independently to down-regulate the pro-inflammatory cytokine TNF-a in human placental trophoblast cells [116-118]. Even the TNF-a has been associated with infection-mediated preterm birth [119, 120]. These studies have suggested that the administra- tion of probiotics may help restoring and maintaining a healthy vaginal microbiota, and thus, that some pro- biotic strains seem to have the ability to interfere with the inflammatory pathway leading to premature deliv- ery. Findings such as these lead us to suggest that some kind of immune modulation by the vaginal microbiota may be opposing the exacerbated inflammation pro- cess observed during labor.Now, which is the importance of the vaginal micro- biota for thenewborn?At the time ofbirth, the newborn is exposed to maternal microbiota [121-124]. The passage through the genital tract is an essential part of this process because the mode of delivery alters the neonatal microbiome [121-126]. In this context, Dominguez-Bello et al. have shown that vaginally- delivered newborns acquired bacterial communities resemble their own mother's vaginal microbiota (dom-inated by Lactobacillus, Prevotella, or Sneathia spp.);and that caesarian section newborns harbored bacterial communities that were more similar to those found on the skin surface, dominated by Staphylo-coccus, Corynebacterium, and Propionibacterium spp. [121]. Other results indicate that Lactobacillus found in the maternal vagina influence the development of the neonatal immune system [127]. Maternal antibi- otics during labor were associated with a decreased transmission rate of vaginal Lactobacillus flora to the neonate during birth, which may have a preventive role in the development of allergic diseases in the newborn [128]. This evidence could be supported by the find- ings of Reid et al., because they have found that in some cases of bacterial infections, nonspecific antibi- otics are used, killing the non-pathogenic members of the microbiota as well as the pathogens and leading to a substantial delay in the restoration of a healthy microbiota [129].EVOLUTIONARY PERSPECTIVESAs in humans, several taxa have developed associ- ations with microbial life, affecting different aspects of their health, sociality and reproduction [e.g., insects 130, 131; aquatic organisms 132; amphibians 133; reptiles 134-136; birds 137, 138; carnivores 139, 140, primates 141, 142]. Accordingly, the vaginal micro- biota may also be regarded as a symbiotic relationship. Symbioses are believed to be ancient, as they involve a long history of interdependence and natural selection involving the coevolution of microbial and host traits that reinforce the relationship [143].Different species use a variety of behaviors which, acting also as social bonding mechanisms, may allow for the direct and indirect transmission of microbiota [144], including licking [145] and grooming [146]. For instance, crucial to regulation of mammalian develop- ment are maternal biochemicals provided to offspring through lactation [147]: an ancient adaptation aris- ing several million years ago, with the function of protecting the product from both desiccation and col- onization by microbial pathogens [148]. A study by Bailey and Coe [149] signals the importance of the mother-infant relationship. These authors found that in infant rhesus monkeys, the integrity of their micro- biota suffered important modifications after separation from their mothers, suggesting that this social disrup- tion lead to an internal environment that was more prone to infection by pathogens. Indeed, it is during suckling when infants receive specific microbe strains from their mothers [150]. A variety of milk compo- nents nourish a newborn by providing the compounds that promote growth, modulate the immune system, promote cognitive development and help establishing a normal gastrointestinal microbiota [151]. For example, human milk contains large amounts of oligosaccha- rides that are not directly assimilated by infants. Instead, these oligosaccharides feed the bacteria B. longum infantis, a primary link in the process of infantís nutrition that helps modulating the immune responses in the intestines and with an important role in the bioconversion of digested nutrients [152]. Some authors have suggested that, compared to that found in other primates and probably due to their beneficial microbiological properties, the significantly greater oligosaccharide concentration found in human milk could have been selected based on the particular dis- ease ecology of humans [153].Recent developments in metagenomic methods allow the identification of the microbial communities composition by means of DNA sequencing, without the need for specific cultures, providing important insights into within and inter-individual variation [154]. Evidence suggests that, in contrast to bacterial commu- nities found in non-human primates and as described before, the healthy human vaginal microbiota is char- acteristically dominated by the presence of Lactobacil- lus [30]. Such findings allow asking about the selective pressures leading to the establishment of this relation- ship across the evolutionary history of the organisms involved. Stumpf et al. [30] have recently suggested three hypotheses explaining the comparative unique- ness of the human vaginal microbiome. Their first argument is that, given that the abundant Lactobacillus are related to estrogen levels in women of reproductive age, their presence could be related to differences in the reproductive cycles between humans and non-human primates. As the vaginal microbiota of most other non- human primates of reproductive age resembles that of non-reproductive human females, its presence could offer protective advantages from pathogens during the crucial periovulatory periods [30]. In this sense, com- parative evidence from apes suggests that menopause is not common among chimpanzees [155], gorillas [156] or orangutans [157], and therefore, that the abun- dant presence of Lactobacillus and menopause could be both derived features of the human life-cycle [155,16992602211070MotherMother43294302325370Dominance of the Lactobacillus species: L. iners L. críspatus L. jensenii L. johnson i iDominance of the Lactobacillus species: L. iners L. críspatus L. jensenii L. johnson i i53841651680845=3Onc_QJ¡zr.o=3O)=3Onc_QJ¡zr.o=3O)53841652557145O"3-’O"3-’20224753215640SymbiosisSymbiosis31349952891155VaginalmicrobiotaVaginalmicrobiotaPregnancyActive immune tolerance► Anti-inflammatory—MjiÁwvresponse♦*’■.Heart Rate Fluctuations19462751410970FetusFig. 3. Suggested interaction among mother, fetus and vaginal microbiota during labor. A sterile anti-inflammatory response is manifested during low-risk labor at term, by either the action of a cholinergic pathway, uterine-like myokines or vaginal microbiome (Image adapted from [159]).158]. The second hypothesis of Stumpf et al. is that the uniqueness of the human vaginal microbiome may be related to human's unique sexuality, i.e., the continuous female sexual receptivity increases females exposure to greater risks of infection and thus a symbiosis that could, somehow, minimize such risks would be highly advantageous and selected. Perhaps current devel- opments involving the primate vaginal microbiome should focus research efforts on the wide variety of primate's reproductive systems and expand our knowl- edge on this aspect of the distinctiveness of the human primate [30]. Last but not least, by means of an “obstet- ric protection hypothesis”, Stumpf et al. suggest that the size of the product in relation to the size of the pelvic outlet could increase the risks of infection to the mother and the fetus, and therefore that a symbiosis reducing such risks would be highly advantageous [30].CONCLUSIONOn the basis of this review, we consider low- risk labor at term as a sterile inflammatory event where three important entities come into interplay (Fig. 3). The first is the mother, whose HRF anal- ysis provides a non-invasive tool to disentangle the role of the anti-inflammatory vagal reflex during labor. The second entity is the vaginal microbiota, where its immunomodulation properties may be based on the modification of bacterial colonies; in particular, the family of Lactobacilli seems to play a key role in this process. The final entity is the fetus, play- ing an important role on the labor process; the fetal heart rate fluctuations may also represent another important marker to understand more about the anti- inflammatory process during the process of labor. While this is a promising path for future research, in this review we have focused our efforts in underlying the primary role of the mother.ACKNOWLEDGMENTSThe support of volunteer mothers and staff of CIMIGen is gratefully acknowledged. J. Javier Reyes- Lagos thanks the Mexican Council for Science andTechnology (CONACyT) for providing fellowship (CVU/Fellowship number: 381983/253449).CONFLICT OF INTERESTAuthors report no conflict of interest.REFERENCESGolightly E, Jabbour HN, Norman JE. Endocrine immune interactions in human parturition. Mol Cell Endocrinol. 2011; 335(1): 52-59.Norwitz ER, Robinson JN, Challis JR. The control of labor. N Engl J Med. 1999; 341(9): 660-666.Peltier MR. Immunology of term and preterm labor. 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Am J Phys Anthropol. 2006; 131: 511521.Wich SA, Utami-Atmoko S, Mitra Setia T, Rijksen H, Schur- mann,etal.LifehistoryofwildSumatranorangutans(Pongo abelii). J Hum Evol. 2004; 47: 385-398.Hawkes K, O'connell JF, Blurton Jones NG, Alvarez H, Charnov EL. Grandmothering, menopause, and the evolu- tion of human life histories. 1998; 95: 1336-1339.Seeley R, Stephens T, Tate P. Anatomy and Physiology. 7th Edition. San Francisco: McGraw-Hill, 2006.20866101230630J.J. Reyes-Lagos et al. / Does Labor Involve a Sterile Anti-Inflammatory Process? PAGE \* MERGEFORMAT #J.J. Reyes-Lagos et al. / Does Labor Involve a Sterile Anti-Inflammatory Process? PAGE \* MERGEFORMAT #8616951230630 PAGE \* MERGEFORMAT #J.J. Reyes-Lagos et al. / Does Labor Involve a Sterile Anti-Inflammatory Process? PAGE \* MERGEFORMAT #J.J. Reyes-Lagos et al. / Does Labor Involve a Sterile Anti-Inflammatory Process?Corréspondéncé to: Dr. G. Pacheco-Lopez, UAM-Lerma, Av. delas Garzas 10, Col. El Panteoín, Lerma, Edo Mexico, C.P. 52005, México. 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