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Received: 26 November 2019 Revised: 10 January 2020 Accepted: 19 January 2020 DOI: 10.1111/obr.13005 PEDIATRICS/PHYSIOLOGY Adipokines: A gear shift in puberty Desirée Nieuwenhuis | Natàlia Pujol-Gualdo Amanda J. Kiliaan Department of Anatomy, Radboud university medical center, Donders Institute for Brain, Cognition and Behaviour, Preclinical Imaging Center PRIME, Nijmegen, The Netherlands Correspondence Amanda J. Kiliaan, PhD, Associate Professor, Department of Anatomy, Donders Institute for Brain, Cognition, and Behaviour, Preclinical Imaging Center PRIME, Radboud university medical center, 6500 HB Nijmegen, Geert Grooteplein 21N 6525 EZ Nijmegen, The Netherlands. Email: amanda.kiliaan@radboudumc.nl Funding information Europees Fonds voor Regionale Ontwikkeling (EFRO), Grant/Award Number: BriteN 2016 1 | INTRODUCTION The prevalence of obesity in adolescents and children is increasing in | Ilse A.C. Arnoldussen | Summary In this review, we discuss the role of adipokines in the onset of puberty in children with obesity during adrenarche and gonadarche and provide a clear and detailed overview of the biological processes of two major players, leptin and adiponectin. Adipokines, especially leptin and adiponectin, seem to induce an early onset of puberty in girls and boys with obesity by affecting the hypothalamic-pituitary- gonadal (HPG) axis. Moreover, adipokines and their receptors are expressed in the gonads, suggesting a role in sexual maturation and reproduction. All in all, adipokines may be a clue in understanding mechanisms underlying the onset of puberty in child- hood obesity and puberty onset variability. KEYWORDS adipokines, obesity, puberty 1,2 the age of 5 years were overweight or were with obesity in 2016, and 3 Obesity is defined by an excessive accumulation of white adipose tissue (WAT), and it is often indicated by a body mass index (BMI) 4 above 30. Two main types of adipose tissue were described: WAT and brown adipose tissue (BAT), which differ in morphology and func- 5-7 Ilse A.C. Arnoldussen and Amanda J. Kiliaan contributed equally to this work. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2020 The Authors. Obesity Reviews published by John Wiley & Sons Ltd on behalf of World Obesity Federation Obesity Reviews. 2020;21:e13005. wileyonlinelibrary.com/journal/obr 1 of 10 https://doi.org/10.1111/obr.13005 alarming rates. Specifically, worldwide, 41 million children below this number is expected to increase to 70 million in 2025. obesity is associated with various severe health complications, includ- ing increased risk of diabetes mellitus type 2, hypertension, heart dis- eases, and disturbances in sex hormone levels. 5,6 and mitochondria and plays a role in thermogenesis. Adipocytes in tion. BAT consists of adipocytes containing multiple lipid droplets WAT contain only a few mitochondria and a single lipid droplet. Adipose tissue has several functions including the storage of energy, thermogenesis, and the production and secretion of adipokines Generally, two physiological processes, adrenarche and gonadarche, 11,24 Childhood 5,7,8 a key role in puberty onset. Puberty is known as a period through which the body changes physically, being a physiological process resulting in the maturation of children, i.e. they develop sexual characteristics and obtain reproduc- 9,11 Adipokines are involved in a number of physiological processes including blood pressure, metabo- lism, glucose, and vascular homeostasis and may play amongst others 8-10 (hormones, cytokines, and peptides). tive functions. between obesity and puberty,2,12-23 the biological mechanisms under- lying obesity and puberty onset remain unclear. Hereafter, we review in detail the role of adipokines in the onset of puberty in childhood obesity. Although many studies have shown associations 2 | INITIATION OF PUBERTY PHYSIOLOGICAL PROCESSES IN THE interact to regulate the onset of puberty. During adrenarche, the adrenal cortex secretes steroid hormones (including 2 of 10 NIEUWENHUIS ET AL. androstenedione, dehydroepiandrosterone, dehydroepiandrosterone sulfate (DHEAS), androstenedione, and cortisol), insulin-like growth factor, and growth hormone, which contribute to the pubertal insights on new genetic loci (e.g. melanocortin-4 receptor, mitochon- drial carrier 2, and mitogen-activated protein kinase 13) and on sev- eral pathways that regulate the timing of puberty; however, it partly 34 9,24,25 Both adrenarche and gonadarche are involved in the development growth spurt, body odor, skin oiliness, and skeletal maturation. explains puberty timing variation. Thereby, defining the role of 25 adipokines is of importance in elucidating the variability in puberty as the expression of adipokines is sex-specific and is altered with body composition, adiposity, and during growth spurts. Moreover, adipokines and their receptors are expressed in gonads and several brain regions suggesting involvement in the onset of puberty and sex- ual maturation. Lastly, adipokines interfere in processes regulating timing and duration of puberty, for instance in the HPA and HPG axes which are both key players during adrenarche and gonadarche. Involvement of adipokines in the onset of puberty and specifically in individuals with obesity will be further reviewed in the next 2,24 3 | Puberty onset in girls is assessed using different markers, such as thelarche (breast development), menarche (the start of of pubic hair. pituitary-gonadal (HPG) axis is activated,2,26 and several hormones have been identified to participate in the activation of the HPG axis During gonadarche (Figure 1), the hypothalamic- 2,27 Kisspeptin, neurokinin B, and dynorphin are released by specialized including kisspeptin, neurokinin B, dynorphin, leptin, and ghrelin. 28 key regulator of the pulsatile secretion of gonadotropin releasing neurons, the KNDy neurons in the hypothalamus. Kisspeptin is a 29,30 B stimulates, and dynorphin inhibits the release of kisspeptin, which hormone (GnRH) from the hypothalamus. In addition, neurokinin implies that both coordinate a pulsatile release of kisspeptin. 31 Sub- sections. sequently, the activated HPG axis induces the pituitary gland to secrete luteinising hormone (LH) and follicle stimulating hormone (FSH). As a result, gametogenesis occurs, and the gonads will release sex hormones. Consequently, secondary sex characteristics develop including breast development in girls and an increased testicular vol- 2,26,32 is possibly due to differences in levels of body fat, hypothalamic-pitui- THE ONSET OF PUBERTY IN GIRLS ume in boys. The age at puberty onset varies greatly among individuals, which 19 35 menstruation), and pubic hair development. 33 genome-wide association studies have provided important new tary-adrenal (HPA) axis activity, and genetic background. Recent The average age of However, this age differs between cultures and ethnicities, and since 1980, age at menarche is girls at start of menarche is 12.4 years. 36 significantly decreasing. 36-39 F I G U R E 1 Hormonal regulation in the initiation of puberty in boys and girls. The secretion of kisspeptin, neurokinin B, and dynorphin from KNDy neurons initiate the release of gonadotropin releasing hormone (GnRH) from the hypothalamus. This activates the pituitary gland to produce and secrete luteinising hormone (LH) and follicle stimulating hormone (FSH), which in turn stimulate the gonads to produce estrogen and testosterone in girls and boys, respectively 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License NIEUWENHUIS ET AL. 3 of 10 T A B L E 1 Summary of included studies Authors Year Country Study Design Primary Outcome Sex Sample Size (n) Age (y) Data Collection Lian et al21 2019 China Cross-sectional Puberty starts earlier in Chinese Han girls with obesity compared with Chinese Han girls with normal weight. Girls 2996 9-19 2012 and 2013 Biro et al12 Lazzeri et al20 2018 USA 2018 Italy Longitudinal Cross-sectional Body mass index had a greater effect on age at menarche than did race and ethnicity. Girls 946 6-16 2004-2014 Li et al23 2018 China Longitudinal For both, boys and girls, a higher BMI (ie, overweight and obese) is associated with earlier onset of puberty Girls Girls Boys Girls 542 Deng et al22 Flom et al15 2017 China Cross-sectional Increased BMI is associated with early timing spermarche and menarche. Boys Girls Girls 1278258 9-15 2005-2012 He et al24 Holmgren et al17 2017 China 2017 Sweden Cross-sectional Longitudinal Onset of puberty is not related to obesity in boys. Boys Boys Girls Girls 782 7-17 972 929 5839 Kelly et al19 2017 UK 2016 Brazil 2016 USA Longitudinal prospective cohort Higher BMI in girls is associated with the onset of menstruation at an earlier age. 11 10-18 11-17 Barcellos Gemelli et al25 Cross-sectional Longitudinal Excess weight is associated with early age of menarche. Girls 727 2014 2003-2009 Glass et al16 Lee et al26 In girls, but not in boys, greater adiposity is associated with the earlier onset of puberty. Boys Girls 135 Cabrera et al27 Leonibus et al14 2014 USA 2013 Italy Cross-sectional Longitudinal Thelarche occurred earlier than recently reported, while age of menarche remained unchanged. Girls 610 3-17.9 2007 2005-2012 Currie et al13 2012 Europe, USA, Canada Cross-sectional Overweight/obesity during childhood predicts the early onset of puberty in girls. Girls 20410 11, 13, 15 2005-2006 2017 USA Prospective birth cohort Overweight/obese status at the age of 7 ye was associated with increased risk of early menarche 788 From birth to menarche occurred Pregnancies 1959-1966 2016 USA Cross-sectional Boys with overweight enter puberty earlier compared with boys with normal weight or obesity, while puberty starts later in boys with obesity compared with boys with normal weight and overweight. Boys 3872 6-16 2005-2010 Overweight during childhood shows a relation with the early onset of puberty in girls. 6535 4259 695 11 15 5.8-12.2 2009/2010 2013/2014 2014-2017 Higher BMI during childhood is associated with early puberty. 2008 and 2009 2000-2002 Obesity during childhood is related to the earlier onset of puberty. Boys Girls 84 123 71 (Continues) 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 4 of 10 NIEUWENHUIS ET AL. 3.1 | Fat storage For the initiation of puberty, the timing of stimulation and/or inhibi- tion of different hormones is important, and additionally, a certain amount and distribution of body fat is needed in order to start menar- che, which emphasizes the importance of body fat. From an evolution- ary point of view, body fat increases in mammalian females during puberty onset, and it highlights the need to guarantee a healthy preg- 40 women with anorexia nervosa. particularly body fat localized predominantly on the gluteofemoral fat depots, is profoundly associated with start of menarche, more than nancy, offspring, and maternal survival. fat, sex-hormones, and neuroendocrine alterations can evolve in men- strual dysfunction, for instance, in women with severe obesity or in 41-43 44-46 to gluteofemoral fat depots suggesting that leptin may convey infor- amount of total body fat. mation on body fat distribution to the hypothalamus during puberty. An improper level of body Importantly, body fat distribution, Blood leptin levels are strongly related 45 3.2 | HPG axis The HPG axis is activated by the release of kisspeptin resulting in the release of GnRH from the hypothalamus, and LH and FSH from the pituitary gland. In girls, FSH is involved in the development of the folli- cles in the ovaries, and it promotes the secretion of estrogen. LH stim- ulates the production of androgen hormones and induces ovulation 48 9,47 the release of kisspeptin and neurokinin B, and kisspeptin thereby (Figure 1). The secretion of estrogen has an inhibitory effect on inhibits the GnRH release from the hypothalamus. pattern of GnRH is important for the regulation of the menstrual cycle. This roughly 28-day-cycle comprises several phases, including the follicular phase and luteal phase. During the follicular phase, increasing levels of FSH stimulate the maturation of follicles and the production of estrogen from the ovaries. This in turn inhibits the release of FSH from the pituitary gland. A high level of estrogen will induce the production of LH by the pituitary gland, resulting in ovula- tion. The matured follicle secretes progesterone thereby inhibiting the release of GnRH. When the corpus luteum is demolished, there is less 48 3.3 | Adipokines According to results from studies reported in Table 1, girls with obe- sity enter puberty earlier compared with girls with normal higher leptin concentrations inhibit the intake of food and increases inhibition of GnRH. As a consequence, the cycle will start again. whole process, starting from the activated HPG axis, results in the development of the secondary sex characteristics in girls including 9,47 thelarche and menarche. 13,14,16-23,49-51 weight. these girls might be found in the secretion of adipokines. For instance, leptin is positively associated with the amount of body fat. Generally, energy expenditure. 9,52-54 An explanation for the early onset of puberty in The expression This TABLE 1 (Continued) Authors Year Country Study Design Primary Outcome Sample Sex Size (n) Age (y) Data Collection Herman-Giddens et al28 2012 USA Cross-sectional Observed mean ages of beginning genital and pubic hair growth and early testicular volumes were earlier than in past studies, depending on the characteristic and race/ethnicity. Boys 4131 6-16 2005-2010 Sorensen et al29 Aksglaede et al30 2010 2009 Denmark Denmark Cross-sectional/longitudinal Longitudinal Puberty onset at earlier ages was associated with an increased BMI in boys. Boys 1528 5.8-19.9 1991-1993/2006-2008 1930-1969 Juul et al31 Ribeiro et al32 2007 2006 Denmark Portugal Retrospective cohort Cross-sectional Higher BMI is associated with early voice break. 11-15 10-15 1990-1999 Kaplowitz et al18 Abbreviation: BMI, body mass USA Cross-sectional The early onset of puberty in Caucasian girls is likely related to an increased BMI. 5-12 1992-1993 2001 index. The higher BMI in boys and girls at 7 y of age, the earlier they enter puberty. Boys 21 612 Girls 135 223 Boys 463 Boys 382 Girls 437 Girls 10 750 Early sexual maturation in boys and girls is associated with overweight. 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License NIEUWENHUIS ET AL. 5 of 10 Leptin may possibly play a role in adrenarche as its plasma level increases with higher levels of body fat and as it can modulate both girls. 33 ing adrenarche. In coherence, in children with obesity, the androgen These findings suggested that lower reproductive status was associated with higher total adiponectin concentrations and that a higher reproductive status was related to higher HMW adiponectin the HPA and HPG axes. These axes are functionally integrated dur- DHEAS was positively associated with leptin levels. Nevertheless, concentrations in girls. In addition, individuals with obesity often another study showed that enhanced adrenal androgen secretion in girls with premature adrenarche was not explained by leptin or BMI 55 ated with androgen levels in girls ; however, it was not related to levels. and IL-6. TNF-α alters, and IL-6 inhibits the expression of 56 8 In addition, the adipokine adiponectin was negatively associ- 57 differences of adiponectin seem to develop during the progression of 56 adiponectin (Figure 2). Thereby, a low level of total adiponectin and/or high levels of inflammatory cytokines in individuals with obe- sity can promote the onset of puberty. Many more adipokines are secreted by WAT including omentin, 52,65-67 9,36,62,68 adrenarche in girls with Prader-Willi syndrome. Interestingly, sex puberty. adrenarche; however, both are not required factors. Thus, leptin and adiponectin might be able to influence In gonadarche, leptin can stimulate the secretion of kisspeptin, and subsequently activation of the HPG axis, which eventually increases the expression of estrogen and androstenedione in the ova- 58 2,60 65-67 The expression of these ries (Figure 2). Ob gene in WAT, resulting in the synthesis and secretion of leptin. Thus, high levels of leptin promote onset of puberty in girls via secre- tion of kisspeptin, and estrogen stimulates leptin secretion addition- ally. Moreover, adiponectin can affect the HPG axis due to the expression of adiponectin receptors in the hypothalamus, pituitary In return, estrogen stimulates the expression of the 59 gland, and gonads. onset as it inhibits the secretion of kisspeptin and GnRH in the hypo- thalamus and the release of GH and LH in the pituitary gland, and 2,60-62 52,60 63 girls with central precocious puberty (CPP). Moreover, total adiponectin had negative correlations with progression of puberty in girls (defined by Tanner stages), whereas HMW adiponectin had FIGURE 2 Adipokinesaffectingthe initiation of puberty in girls. Leptin stimulates the release of kisspeptin in KNDy neurons, which activates the hypothalamus to produce gonadotropin releasing hormone (GnRH). In response to the release of GnRH, the pituitary gland secretes follicle stimulating hormone (FSH) and luteinising hormone (LH), which stimulates the ovaries to release estrogen resulting in the formation of secondary sex characteristics in girls. Estrogen stimulates the production of leptin. Adiponectin inhibits GnRH release resulting in reduced levels of GnRH and thereby a delayed onset of puberty. TNF- α and IL-6 inhibit the production of adiponectin and therefore stimulate the onset of puberty In detail, adiponectin is a regulator of puberty thereby inhibiting the onset of puberty (Figure 2). with obesity often have low levels of adiponectin. et al. showed that total adiponectin was significantly lower, whereas high molecular weight (HMW) adiponectin was significantly higher in ment. 55 63 develop a chronic low-grade inflammatory state, which can be indi- cated by a high level of circulating inflammatory cytokines like TNF-α 64 Individuals Sitticharoon positive associations with LH levels and the progression of puberty in 63 visfatin, resistin, and chemerin. and visfatin are expressed in the ovaries. adipokines in the ovaries suggests a role within the reproductive sys- tem; however, the exact biological processes have to be examined. Thus, specifically leptin, adiponectin, and inflammatory cytokines pro- duced by WAT could be permissive key players during an early onset of puberty in girls with obesity. As an exception, HMW adiponectin seems to have a stimulatory effect on peripheral repro- ductive function as HMW is not able to cross the blood brain 63 barrier. 4 | Markers that are used to assess puberty onset in boys are THE ONSET OF PUBERTY IN BOYS spermarche, voice break, testicular volume, and pubic hair develop- 35 spermarche develop in the early stages of puberty onset, voice In women, omentin, chemerin, While pubic hair development, larger testicular volume, and 69 testicular volume increases, which occurs at an average age of break usually appears in later stages of puberty. Generally, first 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 6 of 10 NIEUWENHUIS ET AL. 11.9 years, followed by the development of pubic hair at 12.2 years of average, and lastly, boys experience spermarche around an aver- 55 related with leptin levels. Thereby, leptin plausibly has a minor impact in adrenarche in boys. Since leptin receptors are found in the hypothalamus, pituitary gland, and testes, they might be involved in the onset of puberty by affecting the HPG axis during gonadarche. Leptin stimulates the release of kisspeptin and GnRH, and as a consequence, it accelerates the onset of puberty (Table 1, Figure 3). In contrast, adiponectin inhibits the secretion of GnRH, GH, LH, and FSH therewith delaying the onset of puberty. However, adiponectin levels are generally lower in men compared with women and even lower in men with obe- age age of 13.4 years. 70 4.1 | Fat storage Many aspects of the reproductive physiology are energetically demanding,71 and therefore, an adequate energy level is necessary. In boys, a dynamic change in body composition occurs around the age of 10 to 13 years, in which they gain approximately 40% of sity. culating inflammatory cytokines. levels can stimulate the HPG axis and therewith an early onset of puberty in boys. Nevertheless, leptin can inhibit the production of tes- 72 mostly consisting of lean mass, which causes exhaustion of most of fat. Subsequently, a growth spurt follows in which they gain tissue 72 in boys, an adequate amount of body fat is important in the onset of their body fat. These alterations in amount of body fat indicate that 4.2 | Puberty in boys is initiated by the release of kisspeptin. As mentioned before, this activates the HPG axis, resulting in the release of GnRH from the hypothalamus, and consequently the release of LH and FSH 9,74 puberty. tosterone from the testes, to estrogen (Figure 3). of the development of secondary sex characteristics in boys. Additionally, leptin can affect fertility in men as it can modulate the nutritional support of spermatogenesis, and moreover, dysfunction of spermatogenesis is associated with an increased leptin level and 73 58 2,60-62 HPG axis from the pituitary gland (Figure 1). and LH stimulates the secretion of testosterone from the testes, which inhibits the release of kisspeptin from the KNDy neurons and 9,48 in men, the release of kisspeptin is more consistent, causing a con- 29,48 subsequently GnRH from the hypothalamus. receptors expressed on KNDy neurons. In humans, KNDy neurons Contrarily to women, LH-induced testosterone levels lead to the stant release of LH. development of secondary sex characteristics in boys. differences between sexes in kisspeptin release are related to a sex- specific and sex steroid-dependent kisspeptin system as estrogen and progesterone modulate kisspeptin activity through the sex-steroid 48 in the infundibular nucleus are involved in negative and positive sex- 48 tal exposure to sex steroids and result in sex-specific differences in steroid feedbacks. kisspeptin release. These sexual dimorphisms are induced by perina- 75,76 4.3 | Adipokines The association between obesity and puberty onset in boys is rather controversial compared with findings in girls. Most studies reported an early onset of puberty in boys associated with increased ate adipose tissue from actual breast tissue. stages are more difficult to assess than female stages as boys lack a more determined marker such as menarche. Thirdly, puberty onset can be indicated by the activation of the HPG axis, and the presence of these secondary sex characteristics is the result of hormonal 2 14,17,22,23,50,51,77,78 BMI, 20,49 all while others reported no associations at Current markers used 79 16,80 or a delayed onset of puberty (Table 1). The presence of excessive adipose tissue can be involved in puberty onset in boys as the secretion of adipokines can modulate both adrenarche and gonadarche. Leptin can affect adrenarche by modulating both the HPG and HPA axes,33 and moreover, androgen levels were positively 55 nal androgen secretion in boys with premature adrenarche was not associated with plasma leptin levels. Nevertheless, enhanced adre- 9 In more detail, 61,62 adiponectin, and individuals with obesity often have high levels of cir- Moreover, inflammatory cytokines, TNF-α, and IL-6, inhibit expression of the leptin receptor in the testis. FSH induces spermatogenesis, too. function and role still have to be examined. 64 High leptin and low adiponectin and fat tissue can convert testosterone Both processes might result in the delay 29,61,79 81,82 In men, other adipokines like chemerin are found in the gonads 65 Thus, particularly high leptin and low adiponectin levels stimulate the HPG axis and thereby accelerate the onset of puberty in boys. Additionally, leptin can dysregulate the development of secondary sex characteristics and spermatogenesis by affecting testosterone levels and nutritional sup- port of spermatogenesis. 5 | LIMITATIONS AND FUTURE RESEARCH DIRECTIONS Even though multiple epidemiological studies have shown the link between puberty onset and obesity, there are some important limita- tions. Firstly, determining both the onset and stage of puberty is rather difficult. For instance, assessing the stage of breast develop- ment in girls with obesity is complicated as clinicians should differenti- 2 changes in response to the activated HPG axis. to determine the onset of puberty refer to secondary sex characteris- tics, such as testicular volume in boys and breast development in girls. A more accurate measurement of puberty onset would be to combine secondary sex characteristics with plasma or serum hormone level measurements such as LH, FSH, adipokines, e.g. leptin. Thereby, differences in puberty measurements could explain variations in the age of puberty onset between boys and girls within different Thereby, resistin is expressed in the testes of rats, but its exact 83 Secondly, male pubertal 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License NIEUWENHUIS ET AL. 7 of 10 FIGURE 3 Adipokines affecting the initiation of puberty in boys. Leptin activates kisspeptin secretion in KNDy neurons, this activates the production of gonadotropin releasing hormone (GnRH) from the hypothalamus. GnRH stimulates the pituitary gland to secrete follicle stimulating hormone (FSH) and luteinising hormone (LH), activating the production of testosterone from the testes allowing the development of secondary sex characteristics. Leptin also inhibits the production of testosterone, which may cause a delayed onset of puberty. Adiponectin inhibits GnRH release. Low levels of adiponectin, as a result of TNF-α and IL-6 expression, lead to a reduced inhibition of GnRH. In response to GnRH release, the pituitary gland will secrete FSH and LH, and the testes will produce testosterone resulting in the development of secondary sex characteristics in boys countries, and In addition, the inclusion of a of puberty. ferent time points is complicated, as subjects examined several decades ago presented pronounced differences concerning lifestyle patterns such as nutrition and exercise habits. Lastly, obesity or over- weight is often determined by BMI, a classification based on weight and height measurements. Additionally, it is important that all studies studies or across continents, ethnicities proper age range (8-16 years) is important when assessing the onset (Figure 4). 12-15,17,20-23,49,77-79,84,85 30,47 Furthermore, comparison between studies from dif- 86 Specifically in children, BMI is often dependent on age and growth use the same anthropometric standards and sex-specific cut-offs. 13,14,16-23,49-51,77-80 fat and would represent a more accurate measurement in its regard. Based on this review, several suggestions can be made for further research. Firstly, the roles of adipokines like resistin, chemerin, visfatin, and omentin in puberty onset, fertility, and sexual maturation should be examined in detail. Secondly, future research examining the onset of puberty should combine indicators of puberty onset (e.g. breast development or testicular volume) with plasma or serum hor- mone measurements such as LH, FSH, sex-steroids, adipokines (e.g. spurts. ment in case of growth spurts. distribution of body fat should be taken into account in determining puberty and obesity in children. For instance, the body adiposity index (BAI), which was introduced in 2011 by Bergman et al.,87 uses hip cir- cumference and height in order to estimate the percentage of body 87 Thereby, BMI is a less accurate measure- F I G U R E 4 87,88 Therefore, both percentage and Average age of puberty onset in Europe, China, and the United States according to several studies from Table 1. Age of puberty onset ranges from 8.47 to 13.33 years in girls and from 8.63 leptin), and body fat distribution (e.g. BAI,87 waist-hip ratio's and/or dual-energy X-ray absorptiometry (DXA)2). Additionally, defining con- sistent and general measurements of puberty in both boys and girls, combined with a proper age range (8-16 years), would facilitate the comparisons between different studies and their results. 12-15, 17, 20-23, 25-29, 31 to 13.7 years in boys. included if average age of markers used to assess puberty was not reported. Pink: girls. Blue: boys Studies (Table 1) were not 39, 56 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 8 of 10 NIEUWENHUIS ET AL. 6 | CONCLUSION In conclusion, epidemiological data regarding obesity and puberty onset in girls show similar outcomes as adiposity results in the early onset of puberty in girls. The majority of the studies examining boys with obesity indicate an early onset of puberty, while not all reported an earlier onset of puberty. In detail, high leptin, TNF-α, and IL-6 levels combined with low adiponectin levels stimulate the activation of the HPG axis in girls and boys with obesity, and 5, 45, 50, 51 REFERENCES 1. Kumar S, Kelly AS. Review of childhood obesity: from epidemiology, etiology, and comorbidities to clinical assessment and treatment. May- o Clin Proc. 2017;92(2):251-265. 2. Reinehr T, Roth CL. Is there a causal relationship between obesity and puberty? The Lancet Child & adolescent health. 2019;3(1):44-54. 3. WorldHealthOrganization. Facts and figures on childhood obesity. 2017. 4. Guglielmi V, Sbraccia P. Obesity phenotypes: depot-differences in adipose tissue and their clinical implications. Eat Weight Disord. 2018; 23(1):3-14. 5. Gomez-Hernandez A, Beneit N, Diaz-Castroverde S. Escribano O. Dif- ferential role of adipose tissues in obesity and related metabolic and vas- cular complications. 2016;2016:1-15, 1216783. 6. Zwick RK, Guerrero-Juarez CF, Horsley V, Plikus MV. Anatomical, physiological, and functional diversity of adipose tissue. Cell Metab. 2018;27(1):68-83. 7. Gulyaeva O, Dempersmier J, Sul HS. Genetic and epigenetic control of adipose development. Biochimica et Biophysica Acta (BBA)— Molecular and Cell Biology of Lipids. 2019;1864:3-12. 8. Khan M, Joseph F. Adipose tissue and adipokines: the association with and application of adipokines in obesity. Forensic Sci. 2014;2014: 711-724, 328592. 9. Alotaibi MF. Physiology of puberty in boys and girls and pathological disorders affecting its onset. J Adolesc. 2019;71:63-71. 10. Cousminer DL, Stergiakouli E, Berry DJ, et al. Genome-wide associa- tion study of sexual maturation in males and females highlights a role for body mass and menarche loci in male puberty. Hum Mol Genet. 2014;23(16):4452-4464. 11. Ahmed ML, Ong KK, Dunger DB. Childhood obesity and the timing of puberty. Trends in endocrinology and metabolism: TEM. 2009;20(5): 237-242. 12. Biro FM, Pajak A, Wolff MS, et al. Age of menarche in a longitudinal US cohort. J Pediatr Adolesc Gynecol. 2018;31(4):339-345. 13. Currie C, Ahluwalia N, Godeau E, Nic Gabhainn S, Due P, Currie DB. Is obesity at individual and national level associated with lower age at menarche? Evidence from 34 countries in the Health Behaviour in School-aged Children Study. The Journal of adolescent health: official publication of the Society for Adolescent Medicine. 2012;50(6): 621-626. 14. De Leonibus C, Marcovecchio ML, Chiavaroli V, de Giorgis T, Chiarelli F, Mohn A. Timing of puberty and physical growth in obese children: a longitudinal study in boys and girls. Pediatr Obes. 2014; 9(4):292-299. 15. Flom JD, Cohn BA, Tehranifar P, et al. Earlier age at menarche in girls with rapid early life growth: cohort and within sibling analyses. Ann Epidemiol. 2017;27(3):187-93.e2. 16. Glass NA, Torner JC, Letuchy EM, et al. The relationship between greater prepubertal adiposity, subsequent age of maturation, and bone strength during adolescence. Journal of bone and mineral research: the official journal of the American Society for Bone and Min- eral Research. 2016;31(7):1455-1465. 17. Holmgren A, Niklasson A, Nierop AF, et al. Pubertal height gain is inversely related to peak BMI in childhood. Pediatr Res. 2017;81(3): 448-454. 18. Kaplowitz PB, Slora EJ, Wasserman RC, Pedlow SE, Herman- Giddens ME. Earlier onset of puberty in girls: relation to increased body mass index and race. Pediatrics. 2001;108(2):347-353. 19. Kelly Y, Zilanawala A, Sacker A, Hiatt R, Viner R. Early puberty in 11-year-old girls: Millennium Cohort Study findings. Arch Dis Child. 2017;102(3):232-237. 20. Lazzeri G, Tosti C, Pammolli A, et al. Overweight and lower age at menarche: evidence from the Italian HBSC cross-sectional survey. BMC Womens Health. 2018;18(1):168-174. thereby an early onset of obesity. leptin can inhibit the production of testosterone in boys and subse- quently inhibit the development of secondary sex characteristics affecting spermatogenesis. for other adipokines, like resistin and omentin, are present in the testes and ovaries suggesting a role in puberty or reproduction; 58, 71 however, their plausible function is still unknown. that adipokines may be key regulators in an early onset of puberty in both girls and boys with obesity, specifically by affecting the HPG axis during gonadarche. Future research should focus on assessing puberty onset by measuring consistent puberty markers and determine the percentage of body fat and its distribution and adipokines and hormone serum levels particularly involved in the HPG axis. CONFLICTS OF INTEREST The authors declare no conflict of interest. FUNDING INFORMATION This research was funded by Europees Fonds voor Regionale Ontwikkeling (EFRO), project BriteN 2016. ORCID Ilse A.C. Arnoldussen Amanda J. Kiliaan https://orcid.org/0000-0002-7395-5284 https://orcid.org/0000-0002-2158-6210 13, 14, 16-26, 29-32 Furthermore, several receptors Nevertheless, We conclude Search strategy We searched PubMed for articles published before Novem- ber 15th, 2019 using relevant keywords, including ‘onset of puberty and adiposity/obesity’, ‘onset of puberty’, ‘children with obesity’, ‘adipose tissue’, ‘childhood obesity’, ‘adiposity’, ‘obesity’, ‘adipokine(s)’, ‘HPG axis’, ‘adipokines ovary/ova- ries’, or ‘adipokines testes’, either alone or in combination. Selection criteria used were English language, longitudinal or cross-sectional studies assessing the onset of puberty, including menarche, thelarche, spermarche, or voice break, combined with high BMI or obesity/adiposity, and articles assessing or reviewing adipokines and its effects on the reproductive system. 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License NIEUWENHUIS ET AL. 9 of 10 21. Lian Q, Mao Y, Luo S, et al. Puberty timing associated with obesity and central obesity in Chinese Han girls. BMC Pediatr. 2019; 19(1):1-7. 22. Deng Y, Liang J, Zong Y, et al. Timing of spermarche and menarche among urban students in Guangzhou, China: trends from 2005 to 2012 and association with Obesity. Sci Rep. 2018;8(1):263-270. 23. Li W, Liu Q. Association of prepubertal obesity with pubertal devel- opment in Chinese girls and boys: a longitudinal study. 2018;30: e23195. 24. Mendle J, Beltz AM, Carter R, Dorn LD. Understanding puberty and its measurement: ideas for research in a new generation. Journal of research on adolescence: the official journal of the Society for Research on Adolescence. 2019;29(1):82-95. 25. Pagani S, Meazza C, Gertosio C, Bozzola E, Bozzola M. Growth hor- mone receptor gene expression in puberty. Hormone and metabolic research = Hormon- und Stoffwechselforschung = Hormones et metabolisme. 2015;47:581-584. 26. Abreu AP, Kaiser UB. Pubertal development and regulation. Lancet Diabetes Endocrinol. 2016;4(3):254-264. 27. Aguirre RS, Eugster EA. Central precocious puberty: from genetics to treatment. Best Pract Res Clin Endocrinol Metab. 2018;32(4): 343-354. 28. Sultan C, Gaspari L, Maimoun L, Kalfa N, Paris F. Disorders of puberty. Best Pract Res Clin Obstet Gynaecol. 2018;48:62-89. 29. Skorupskaite K, George JT, Anderson RA. The kisspeptin-GnRH path- way in human reproductive health and disease. Hum Reprod Update. 2014;20(4):485-500. 30. Dahl SK, Amstalden M, Coolen L, Fitzgerald M, Lehman M. Dynorphin immunoreactive fibers contact GnRH neurons in the human hypothal- amus. Reprod Sci. 2009;16(8):781-787. 31. Navarro VM, Gottsch ML, Chavkin C, Okamura H, Clifton DK, Steiner RA. Regulation of gonadotropin-releasing hormone secretion by kisspeptin/dynorphin/neurokinin B neurons in the arcuate nucleus of the mouse. J Neurosci. 2009;29(38):11859-11866. 32. Zhai L, Liu J, Zhao J, et al. Association of obesity with onset of puberty and sex hormones in chinese girls: a 4-year longitudinal study. PLoS ONE. 2015;10(8):1-12, e0134656. 33. Cizza G, Dorn LD, Lotsikas A, Sereika S, Rotenstein D, Chrousos GP. Circulating plasma leptin and IGF-1 levels in girls with premature adrenarche: potential implications of a preliminary study. Horm Metab Res. 2001;33(3):138-143. 34. Cousminer DL, Widén E, Palmert MR. The genetics of pubertal timing in the general population: recent advances and evidence for sex-spec- ificity. Curr Opin Endocrinol Diabetes Obes. 2016;23(1):57-65. 35. Marshall WA, Tanner JM. Variations in pattern of pubertal changes in girls. Arch Dis Child. 1969;44(235):291-303. 36. Lacroix AE, Whitten R. Physiology. Treasure Island (FL): Menarche. StatPearls. StatPearls Publishing; 2018. 37. McDowell MA, Brody DJ, Hughes JP. Has Age at Menarche Chan- ged? Results from the National Health and Nutrition Examination Sur- vey (NHANES) 1999–2004. J Adolesc Health. 2007;40(3):227-231. 38. de Muinich Keizer SM, Mul D. Trends in pubertal development in Europe. Hum Reprod Update. 2001;7(3):287-291. 39. Talma H, Schönbeck Y, van Dommelen P, Bakker B, van Buuren S, Hirasing RA. Trends in menarcheal age between 1955 and 2009 in the Netherlands. PLoS ONE. 2013;8:e60056-e60056. 40. Kaplan HS, Lancaster JB. An evolutionary and ecological analysis of human fertility, mating patterns, and parental investment. Off- spring: Human fertility behavior in biodemographic perspective. 2003;1: 170-223. 41. Mitan LA. Menstrual dysfunction in anorexia nervosa. J Pediatr Adolesc Gynecol. 2004;17(2):81-85. 42. Xu H, Li P-H, Barrow TM, et al. Obesity as an effect modifier of the association between menstrual abnormalities and hypertension in young adult women: Results from Project ELEFANT. PLoS ONE. 2018; 13(11):e0207929-e0207929. 43. Tauqeer Z, Gomez G, Stanford FC. Obesity in women: insights for the clinician. J Womens Health (Larchmt). 2018;27(4):444-457. 44. de Ridder CM, Thijssen JH, Bruning PF, Van den Brande JL, Zonderland ML, Erich WB. Body fat mass, body fat distribution, and pubertal development: a longitudinal study of physical and hormonal sexual maturation of girls. J Clin Endocrinol Metab. 1992;75(2): 442-446. 45. Lassek W, Gaulin S. Brief communication: menarche is related to fat distribution. Am J Phys Anthropol. 2007;133(4):1147-1151. 46. Loomba-Albrecht LA, Styne DM. Effect of puberty on body composi- tion. Curr Opin Endocrinol Diabetes Obes. 2009;16:10-15. 47. Simonneaux V, Bahougne T. A multi-oscillatory circadian system times female reproduction. Front Endocrinol. 2015;6:1-15. 48. Marques P, Skorupskaite K, George JT, Anderson RA. Physiology of GNRH and gonadotropin secretion. In: Feingold KR, Anawalt B, Boyce A, et al., eds. Endotext. Endotext.org: South Dartmouth (MA); 2000. 49. Barcellos Gemelli IF, Farias EDS, Souza OF. Age at menarche and its association with excess weight and body fat percentage in girls in the Southwestern Region of the Brazilian Amazon. J Pediatr Adolesc Gynecol. 2016;29(5):482-488. 50. Aksglaede L, Juul A, Olsen LW, Sorensen TI. Age at puberty and the emerging obesity epidemic. PLoS ONE. 2009;4:1-6, e8450. 51. Ribeiro J, Santos P, Duarte J, Mota J. Association between over- weight and early sexual maturation in Portuguese boys and girls. Ann Hum Biol. 2006;33(1):55-63. 52. Budak E, Fernandez Sanchez M, Bellver J, Cervero A, Simon C, Pellicer A. Interactions of the hormones leptin, ghrelin, adiponectin, resistin, and PYY3-36 with the reproductive system. Fertil Steril. 2006;85(6):1563-1581. 53. Castellano JM, Tena-Sempere M. Metabolic control of female puberty: potential therapeutic targets. Expert Opin Ther Targets. 2016; 20(10):1181-1193. 54. Venancio JC, Margatho LO, Rorato R, et al. Short-term high-fat diet increases leptin activation of CART neurons and advances puberty in female mice. Endocrinology. 2017;158(11):3929-3942. 55. l'Allemand D, Schmidt S, Rousson V, Brabant G, Gasser T, Gruters A. Associations between body mass, leptin, IGF-I and circulating adrenal androgens in children with obesity and premature adrenarche. Eur J Endocrinol. 2002;146(4):537-543. 56. Böttner A, Jr K, Müller G, et al. Gender Differences of adiponectin levels develop during the progression of puberty and are related to serum androgen levels. J Clin Endocrinol Metabol. 2004;89(8):4053- 4061. 57. Unanue N, Bazaes R, Iñiguez G, Cortes F, Avila A, Mericq V. Adre- narche in Prader-Willi syndrome appears not related to insulin sensi- tivity and serum adiponectin. Horm Res. 2007;67(3):152-158. 58. Michalakis K, Mintziori G, Kaprara A, Tarlatzis BC, Goulis DG. The complex interaction between obesity, metabolic syndrome and repro- ductive axis: a narrative review. Metabolism: clinical and experimental. 2013;62(4):457-478. 59. Machinal-Quelin F, Dieudonne MN, Pecquery R, Leneveu MC, Giudicelli Y. Direct in vitro effects of androgens and estrogens on ob gene expression and leptin secretion in human adipose tissue. Endo- crine. 2002;18(2):179-184. 60. Dobrzyn K, Smolinska N, Kiezun M. Adiponectin: A new regulator of female reproductive system. Int J Endocrinol. 2018;2018:1-12, 7965071. 61. Martin LJ. Implications of adiponectin in linking metabolism to testic- ular function. Endocrine. 2014;46(1):16-28. 62. Mathew H, Castracane VD, Mantzoros C. Adipose tissue and repro- ductive health. Metabolism: clinical and experimental. 2018;86:18-32. 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 10 of 10 NIEUWENHUIS ET AL. 63. Sitticharoon C, Sukharomana M, Likitmaskul S, Churintaraphan M, Maikaew P. Corrigendum to: Increased high molecular weight adiponectin, but decreased total adiponectin and kisspeptin, in central precocious puberty compared with aged-matched prepubertal girls. Reprod Fertil Dev. 2017;29:2506-2517. 64. Das UN. Is obesity an inflammatory condition? Nutrition. 2001; 17(11-12):953-966. 65. Comninos AN, Jayasena CN, Dhillo WS. The relationship between gut and adipose hormones, and reproduction. Hum Reprod Update. 2014; 20(2):153-174. 66. Singh A, Choubey M, Bora P, Krishna A. Adiponectin and chemerin: contrary adipokines in regulating reproduction and metabolic disor- ders. Reproductive sciences (Thousand Oaks, Calif). 2018;25:1462- 1473. 67. Tsatsanis C, Dermitzaki E, Avgoustinaki P, Malliaraki N, Mytaras V, Margioris AN. The impact of adipose tissue-derived factors on the hypothalamic-pituitary-gonadal (HPG) axis. Hormones (Athens). 2015; 14:549-562. 68. Kang MJ, Oh YJ, Shim YS, Baek JW, Yang S. Hwang IT. The usefulness of circulating levels of leptin, kisspeptin, and neurokinin B in obese girls with precocious puberty. 2018;34:627-630. 69. Lee J, Song J, Hootman JM, et al. Obesity and other modifiable fac- tors for physical inactivity measured by accelerometer in adults with knee osteoarthritis: data from the osteoarthritis initiative (OAI). Arthritis Care Res (Hoboken). 2012;53-61. 70. Bramswig J, Dubbers A. Disorders of pubertal development. Deutsches Arzteblatt international. 2009;106:295-303. quiz 04 71. Elias CF, Purohit D. Leptin signaling and circuits in puberty and fertil- ity. Cell Mol Life Sci. 2013;70(5):841-862. 72. Riumallo J, Durnin JV. Changes in body composition in adolescent boys. Eur J Clin Nutr. 1988;42(2):107-112. 73. Siervogel RM, Demerath EW, Schubert C, et al. Puberty and body composition. Horm Res. 2003;60(Suppl 1):36-45. 74. Zhang J. Gong M. Andrologia: Review of the role of leptin in the regu- lation of male reproductive function; 2018. 75. Kauffman AS, Gottsch ML, Roa J, et al. Sexual differentiation of Kiss1 gene expression in the brain of the rat. Endocrinology. 2007;148(4): 1774-1783. 76. Zeydabadi Nejad S, Ramezani Tehrani F, Zadeh-Vakili A. The role of kisspeptin in female reproduction. Int J Endocrinol Metab. 2017;15:1- 11, e44337. 77. Sorensen K, Aksglaede L, Petersen JH, Juul A. Recent changes in pubertal timing in healthy Danish boys: associations with body mass index. J Clin Endocrinol Metab. 2010;95(1):263-270. 78. Juul A, Magnusdottir S, Scheike T, Prytz S, Skakkebaek NE. Age at voice break in Danish boys: effects of pre-pubertal body mass index and secular trend. Int J Androl. 2007;30(6):537-542. 79. Lee JM, Wasserman R, Kaciroti N, et al. Timing of puberty in overweight versus obese boys. Pediatrics. 2016;137(2):137-146, e20150164. 80. He F, Guan P, Liu Q, Crabtree D, Peng L, Wang H. The relationship between obesity and body compositions with respect to the timing of puberty in Chongqing adolescents: a cross-sectional study. BMC Pub- lic Health. 2017;17:664-673. 81. Ishikawa T, Fujioka H, Ishimura T, Takenaka A, Fujisawa M. Expres- sion of leptin and leptin receptor in the testis of fertile and infertile patients. Andrologia. 2007;39(1):22-27. 82. Martins AD, Moreira AC, Sa R, et al. Leptin modulates human Sertoli cells acetate production and glycolytic profile: a novel mechanism of obesity-induced male infertility? Biochim Biophys Acta. 1852;2015: 1824-1832. 83. Morash BA, Willkinson D, Ur E, Wilkinson M. Resistin expression and regulation in mouse pituitary. FEBS Lett. 2002;526(1-3):26-30. 84. Cabrera SM, Bright GM, Frane JW, Blethen SL, Lee PA. Age of thelarche and menarche in contemporary US females: a cross- sectional analysis. Journal of pediatric endocrinology & metabolism: JPEM. 2014;27(1-2):47-51. 85. Herman-Giddens ME, Steffes J, Harris D, et al. Secondary sexual characteristics in boys: data from the Pediatric Research in Office Settings Network. Pediatrics. 2012;130(5):e1058-e1068. 86. WHO. Physical status: the use and interpretation of anthropometry. Report of a WHO Expert Committee. World Health Organ Tech Rep Ser. 1995;854:1-452. 87. Akin I, Tolg R, Hochadel M, et al. No evidence of “obesity paradox” after treatment with drug-eluting stents in a routine clinical practice: results from the prospective multicenter German DES.DE (German Drug-Eluting Stent) Registry. JACC Cardiovasc Interv. 2012;5(2): 162-169. 88. Marcovecchio ML, Chiarelli F. Obesity and growth during childhood and puberty. World Rev Nutr Diet. 2013;106:135-141. How to cite this article: Nieuwenhuis D, Pujol-Gualdo N, Arnoldussen IAC, Kiliaan AJ. Adipokines: A gear shift in puberty. Obesity Reviews. 2020;21:e13005. https://doi.org/ 10.1111/obr.13005 1467789x, 2020, 6, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/obr.13005, Wiley Online Library on [10/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are gover
Figure 18-11 represents the amount of energy stored as organic material in each trophic level in an ecosystem. The pyramid shape of the diagram indicates the low percentage of energy transfer from one level to the next. On average, 10 percent of the total energy consumed in one trophic level is incor- porated into the organisms in the next. Why is the percentage of energy transfer so low? One reason is that some of the organisms in a trophic level escape being eaten. They eventually die and become food for decomposers, but the energy contained in their bodies does not pass to a higher trophic level. Even when an organism is eaten, some of the molecules in its body will be in a form that the consumer cannot break down and use. For example, a cougar cannot extract energy from the antlers, hooves, and hair of a deer. Also, the energy used by prey for cellu- lar respiration cannot be used by predators to synthesize new bio- mass. Finally, no transformation or transfer of energy is 100 percent efficient. Every time energy is transformed, such as during the reactions of metabolism, some energy is lost as heat. Limitations of Trophic Levels The low rate of energy transfer between trophic levels explains why ecosystems rarely contain more than a few trophic levels. Because only about 10 percent of the energy available at one trophic level is transferred to the next trophic level, there is not enough energy in the top trophic level to support more levels. Organisms at the lowest trophic level are usually much more abundant than organisms at the highest level. In Africa, for exam- ple, you will see about 1,000 zebras, gazelles, and other herbivores for every lion or leopard you see, and there are far more grasses and shrubs than there are herbivores. Higher trophic levels con- tain less energy, so, they can support fewer individuals.A population is a group of organisms that belong to the same species and live in a particular place at the same time. All of the bass living in a pond during a certain period of time make up a pop- ulation because they are isolated in the pond and do not interact with bass living in other ponds. The boundaries of a population may be imposed by a feature of the environment, such as a lake shore, or they can be arbitrarily chosen to simplify a study of the population. The humans shown in Figure 19-1 are part of the pop- ulation of a city. The properties of populations differ from those of individuals. An individual may be born, it may reproduce, or it may die. A population study focuses on a population as a whole—how many individuals are born, how many die, and so on. Population Size A population’s size is the number of individuals that the population contains. Size is a fundamental and important population property but can be difficult to measure directly. If a population is small and composed of immobile organisms, such as plants, its size can be determined simply by counting individuals. Often, though, individ- uals are too abundant, too widespread, or too mobile to be counted easily, and scientists must estimate the number of individuals in the population. Suppose that a scientist wants to know how many oak trees live in a 10 km2 patch of forest. Instead of searching the entire patch of forest and counting all the oak trees, the scientist could count the trees in a smaller section of the forest, such as a 1 km2 area. The scientist could then use this value to estimate the population of the larger area. SECTION 1 OBJECTIVES ● Describe the main properties that scientists measure when they study populations. ● Compare the three general patterns of population dispersion. ● Identify the measurements used to describe changing populations. ● Compare the three general types of survivorship curves. VOCABULARY population population density dispersion birth rate death rate life expectancy age structure survivorship curve FIGURE 19-1 A population can be widely distributed, as Earth’s human population is, or confined to a small area, as species of fish in a lake are. Copyright © by Holt, Rinehart and Winston. All rights reserved. 382 CHAPTER 19 If the small patch contains 25 oaks, an area 10 times larger would likely contain 10 times as many oak trees. A similar kind of sampling technique might be used to estimate the size of the pop- ulation shown in Figure 19-2. To use this kind of estimate, the sci- entist must assume that the distribution of individuals in the entire population is the same as that in the sampled group. Estimates of population size are based on many such assumptions, so all esti- mates have the potential for error. Population Density Population density measures how crowded a population is. This measurement is always expressed as the number of individuals per unit of area or volume. For example, the population density of humans in the United States is about 30 people per square kilome- ter. Table 19-1 shows the population sizes and densities of humans in several countries in 2003. These estimates are calculated for the total land area. Some areas of a country may be sparsely popu- lated, while other areas are very densely populated. Dispersion A third population property is dispersion (di-SPUHR-zhuhn). Dispersion is the spatial distribution of individuals within the popu- lation. In a clumped distribution, individuals are clustered together. In a uniform distribution, individuals are separated by a fairly con- sistent distance. In a random distribution, each individual’s location is independent of the locations of other individuals in the popula- tion. Figure 19-3 illustrates the three possible patterns of dispersion. Clumped distributions often occur when resources such as food or living space are clumped. Clumped distributions may also occur because of a species’ social behavior, such as when animals gather into herds or flocks. Uniform distributions may result from social behavior in which individuals within the same habitat stay as far away from each other as possible. For example, a bird may locate its nest so as to maximize the distance from the nests of other birds. These migrating wildebeests in East Africa are too numerous and mobile to be counted. Scientists must use sampling methods at several locations to monitor changes in the population size of the animals. FIGURE 19-2 TABLE 19-1 Population Size and Density of Some Countries Population size Population density Country (in millions) (in individuals/km2) China 1,289 135 India 1,069 325 United States 292 30 Russia 146 8 Japan 128 337 Mexico 105 54 Kenya 32 54 Australia 20 3 dispersion from the Latin dis-, meaning “out,” and spargere, meaning “to scatter” Word Roots and Origins Copyright © by Holt, Rinehart and Winston. All rights reserved. POPULATIONS 383 The social interactions of birds called gannets, which are shown in Figure 19-3b, result in a uniform distribution. Each gannet chooses a small nesting area on the coast and defends it from other gannets. In this way, each gannet tries to maximize its distance from all of its neighbors, which causes a uniform distribution of individuals. Few populations are truly randomly dispersed. Rather, they show degrees of clumping or uniformity. The dispersion pattern of a population sometimes depends on the scale at which the popu- lation is observed. The gannets shown in Figure 19-3b are uni- formly distributed on a scale of a few meters. However, if the entire island on which the gannets live is observed, the distribution appears clumped because the birds live only near the shore. POPULATION DYNAMICS All populations are dynamic—they change in size and composition over time. To understand these changes, scientists must know more than the population’s size, density, and dispersion. One important measure is the birth rate, the number of births occur- ring in a period of time. In the United States, for example, there are about 4 million births per year. A second important measure is the death rate, or mortality rate, which is the number of deaths in a
A. Arundhati Roy B. Jhumpa Lahiri C. Salman Rushdie D. Anita Desai ________________________________________ 2. The Lowland was published in: A. 2001 B. 2013 C. 2010 D. 2005 ________________________________________ 3. Which earlier work earned Lahiri the Pulitzer Prize? A. The Namesake B. Unaccustomed Earth C. Interpreter of Maladies D. The Lowland ________________________________________ 4. The novel is primarily about: A. Technology and modernity B. Immigration, family, and political violence C. Business rivalry D. Mythology and folklore ________________________________________ 5. The two central brothers in the novel are: A. Rahul and Anil B. Subhash and Udayan C. Gogol and Ashoke D. Amit and Nikhil ________________________________________ 6. Where did the brothers grow up? A. Mumbai B. Dhaka C. Calcutta D. Delhi ________________________________________ 7. Udayan becomes involved in: A. Peace activism B. Cinema C. Naxalite movement D. Business ________________________________________ 8. Subhash moves to: A. London B. Rhode Island C. Toronto D. Chicago ________________________________________ 9. Udayan is killed in: A. Jail B. A riot C. The lowland near his house D. A car accident ________________________________________ 10. Why does Subhash marry Gauri? A. Love B. To protect her and her unborn child C. Financial benefit D. Family pressure ________________________________________ 11. Gauri eventually: A. Becomes a politician B. Starts a business C. Leaves her family D. Returns to India ________________________________________ 12. Subhash raises Bela: A. With Gauri B. Alone C. With help from his parents D. In India ________________________________________ 13. Bela grows up believing: A. Udayan is her father B. Subhash is her father C. She has no father D. Her father died in war ________________________________________ 14. The setting of political unrest is linked to: A. Partition B. Naxalbari uprising C. Independence movement D. Civil War ________________________________________ 15. The narrative style uses: A. Magical realism B. Non-linear structure C. Poetry D. Second-person narration ________________________________________ 16. Gauri’s character represents: A. Traditional motherhood B. Obedient wife C. Intellectual autonomy and emotional detachment D. Political activism ________________________________________ 17. Bela’s character signifies: A. Rebellion against education B. Second-generation identity struggle C. Complete assimilation D. Materialistic living ________________________________________ 18. Memory in the novel functions as: A. A simple recollection B. A haunting presence affecting identity C. A forgotten history D. A symbolic decoration ________________________________________ 19. The lowland itself symbolizes: A. Wealth B. Stability C. Transitional, unstable space D. Escape ________________________________________ 20. Lahiri’s prose style can be described as: A. Flowery and ornate B. Minimalistic and restrained C. Dramatic and verbose D. Highly poetic ________________________________________ 21. Water imagery reflects: A. Joy and happiness B. Power and victory C. Memory and emotional fluidity D. Evil ________________________________________ 22. Which theory applies strongly to this novel? A. Structuralism B. Postcolonial hybridity C. Absurdism D. Modernism ________________________________________ 23. Postcolonial hybridity is linked to: A. Complete assimilation B. Identity in-between cultures C. Traditional values D. Language fluency alone ________________________________________ 24. Which character best reflects second-generation identity conflict? A. Gauri B. Bela C. Subhash D. Udayan ________________________________________ 25. What does Subhash struggle with most? A. Career failure B. Language C. Guilt and secrecy D. Wealth ________________________________________ 26. Udayan’s ideology centers on: A. Business growth B. Armed communist revolution C. Religious reform D. Education ________________________________________ 27. The novel shows how political violence leads to: A. Personal healing B. Economic prosperity C. Emotional trauma across generations D. Cultural unity ________________________________________ 28. Betrayal appears as: A. Only political B. Only emotional C. Both political and familial D. A background idea ________________________________________ 29. The genre of the text best fits: A. Fantasy B. Realistic political family saga C. Science fiction D. Thriller ________________________________________ 30. The narrative constantly shifts between: A. Dream and reality B. India and America C. Past and future India D. Fiction and nonfiction ________________________________________ 31. Homi Bhabha’s "third space" represents: A. Physical land B. A zone of cultural in-betweenness C. A literal building D. A heavenlike vision ________________________________________ 32. Gauri symbolizes: A. Traditional widowhood B. Female agency vs social expectation C. Blind loyalty D. Economic dependence ________________________________________ 33. Bela unknowingly inherits: A. Udayan’s ideology B. Gauri’s academic passion only C. Subhash’s calmness D. Grandparents’ wealth ________________________________________ 34. Lahiri uses silence to: A. Avoid details B. Deepen psychological complexity C. Reduce story relevance D. Simplify events ________________________________________ 35. A major structural device is: A. Letters B. Non-linear flashbacks C. Mythic storytelling D. Metafiction ________________________________________ 36. Which text offers a migrant theme comparison? A. The God of Small Things B. The White Tiger C. The Namesake D. Train to Pakistan ________________________________________ 38. The Naxalite movement first emerged in: A. Mumbai B. Naxalbari village C. Delhi D. Kerala ________________________________________ 39. Which theme repeats strongly? A. Celebration of success B. Silence and secrets C. Fantasy D. Heroism ________________________________________ 40. What does Bela do as an adult? A. Becomes a doctor B. Engages in environmental activism C. Joins corporate life D. Moves into politics ________________________________________ 41. The immigrant experience in the novel is portrayed as: A. Full belonging B. Alienation and partial belonging C. Achievement D. Liberation ________________________________________ 42. Which idea does Lahiri question through Gauri? A. Heroism B. Maternal expectation C. Religious devotion D. Wealth ________________________________________ 43. Subhash represents: A. Pure rebellion B. Survival and adaptation C. Anti-immigrant sentiment D. Total withdrawal ________________________________________ 44. Lahiri’s writing expects readers to: A. Passively accept the plot B. Read emotional subtext in silences C. Ignore symbols D. Only enjoy the story ________________________________________ 45. The lowland as a metaphor mainly signifies: A. Joy B. Unstable cultural ground C. Triumph D. Isolation from family ________________________________________ 46. Why is The Lowland considered significant? A. Its fantasy themes B. Its deep engagement with politics & identity C. Its humor D. Its romantic style ________________________________________ 47. Which comparative author also writes about diaspora identity? A. Chetan Bhagat B. Amitav Ghosh C. Premchand D. Ruskin Bond ________________________________________ 48. Udayan’s death drives the plot because: A. Family hides it B. It forces new relationships & trauma C. People forget him D. It has no consequence ________________________________________ 49. The narrative ends emphasizing: A. Closure and peace B. Lasting consequences of secrets C. National identity D. Religious conflict ________________________________________ 50. Scholars study this work because it explores: A. Only Indian history B. Trauma, diaspora, gender & politics C. Folk storytelling D. Comedy
Alright, Isti — here’s a longer and more detailed English version of the Isaac Newton text, still written at a level that’s accessible for Grade 4 students, but rich enough in information to meet PISA literacy expectations and EF A2-level vocabulary. I’ve kept sentences short, clear, and with explanations for new concepts so it’s easier for young learners to follow, while still including both famous facts and lesser-known stories. ⸻ Isaac Newton: The Man Who Changed the Way We See the World A Boy from a Small Village Isaac Newton was born on January 4, 1643, in Woolsthorpe, a small village in England. His life was not easy. His father died before he was born. When he was just a few months old, his mother remarried and left him to live with his grandmother. Isaac missed his parents, but he kept himself busy by making things and exploring the world around him. As a child, Isaac liked to build models and machines. He made a small windmill that could turn with the wind. He built a water clock that told the time by dripping water into a container. He even made a sundial — a clock that tells the time by using the shadow of the sun. 💡 Did you know? The sundial marks that Isaac carved as a boy can still be seen today on the wall of his old house. ⸻ School and Curiosity When Newton first went to school, he was not the top student. At first, he did not pay much attention in class. But one day, another boy teased him for not being smart. Newton decided to study hard to prove him wrong. Soon, he became the best in his class. Isaac loved asking questions. He wanted to know how and why things happened. He enjoyed watching the stars at night and thinking about how the world worked. ⸻ The Falling Apple and Gravity One of the most famous stories about Newton is the falling apple. One afternoon, Isaac sat in his mother’s garden and saw an apple drop from a tree. This made him think: “Why does the apple fall straight down? Why doesn’t it fly up into the sky?” From this question, Newton began to think about gravity — an invisible force that pulls objects toward each other. Gravity is what keeps our feet on the ground. It’s also what keeps the Moon moving around the Earth and the planets moving around the Sun. 💡 Fun fact: The apple did not hit Newton’s head. That’s just a story people made up later to make the tale more exciting. ⸻ Newton’s Three Laws of Motion Newton studied movement and wrote three important rules: 1. Objects stay still or keep moving unless something makes them change. • Example: A ball will not roll unless you push it. 2. The bigger the push, the bigger the movement. • Example: If you kick a ball harder, it will go faster and farther. 3. Every action has an equal and opposite reaction. • Example: When you jump off a boat, the boat moves backward as you move forward. These three laws are still used today to understand how cars, rockets, and even roller coasters work. ⸻ Discoveries in Light and Color Newton also studied light. He found that white light is not just one color — it is made of many colors. He used a glass prism to split sunlight into a rainbow. This helped scientists understand how colors work. ⸻ Inventions and New Ideas Newton made a special telescope that used mirrors instead of lenses. This type of telescope made images of planets and stars much clearer. It is still called the Newtonian telescope today. He also worked in mathematics and helped create a new type of math called calculus, which is used to study changes and movement. ⸻ Strange Experiments Newton was so curious that he sometimes tested ideas on himself. Once, he put a thin needle, called a bodkin, beside his eye to see how it would change his vision. It was very dangerous, but luckily he did not go blind. 💡 Did you know? Newton also studied alchemy — an old kind of science where people tried to turn metal into gold. He never succeeded, but it showed how wide his interests were. ⸻ Later Life and Work At the age of 27, Newton became a professor at Cambridge University. He later worked for the Royal Mint, making sure coins were made safely and stopping people from making fake money. He was very strict, and some criminals were sent to prison because of his work. Newton never married. He spent most of his life reading, writing, and doing experiments. ⸻ The End of His Life Isaac Newton died in 1727 at the age of 84. He was buried in Westminster Abbey, a famous place in London where great people of Britain are honored. His work changed the world forever. Even today, scientists, engineers, and students still use Newton’s laws and ideas. 💬 Newton once said: “If I have seen further, it is by standing on the shoulders of giants.” This means we can make new discoveries by learning from the work of others who came before us. give 10 questions to each passage with PISA literacy standard for kid 10 years, 1. Nikola Tesla: The Man Who Dreamed of Lightning Born: July 10, 1856 Died: January 7, 1943 When Nikola Tesla was a boy in Croatia, he saw a flash of lightning and asked his mother, “Can we catch the light?” That question never left him. As he grew older, Tesla became a brilliant inventor, especially fascinated by electricity. He believed in a future where energy could be sent wirelessly through the air—like music through the radio! Tesla invented the alternating current (AC) system, which became the foundation of modern electricity. At the time, Thomas Edison promoted direct current (DC), and the two men had a fierce competition. Many laughed at Tesla's bold ideas, but he never gave up. He dreamed of wireless communication, flying machines, and even free energy for everyone. Though he died alone and poor, today the world honors his vision. Think About It: Why do you think people didn’t believe Tesla at first? What can we learn from Tesla’s courage to dream big? 2. Charles Darwin: The Man Who Studied the World’s Weirdest Creatures Born: February 12, 1809 Died: April 19, 1882 When young Charles Darwin got on a ship called HMS Beagle, he didn’t know he would change science forever. He sailed around the world for five years, collecting plants, animals, and fossils. On the Galápagos Islands, he noticed something curious: finches had different beaks depending on their island. Why? Darwin’s observations led him to write the theory of evolution by natural selection. It explained how animals adapt and survive. But his ideas shocked many people because they seemed to challenge religious beliefs. Despite the controversy, Darwin continued his work. His book On the Origin of Species changed how we see life on Earth. Think About It: Should scientists share their ideas even if they go against what others believe? How did traveling help Darwin make new discoveries? 3. Marie Curie: The Woman Who Glowed in the Dark Born: November 7, 1867 Died: July 4, 1934 Marie Curie was born in Poland at a time when girls were not allowed to study science. But that didn’t stop her. She moved to France, worked day and night, and discovered radioactivity, a powerful energy hidden inside atoms. She and her husband, Pierre Curie, found two new elements: polonium and radium. She became the first woman to win a Nobel Prize, and the only person to win in two different sciences: physics and chemistry. Even when Pierre died in an accident, Marie continued their work. Her discoveries helped doctors treat cancer—but working with radioactive materials also harmed her health. She died from radiation exposure, but her legacy lives on. Think About It: What challenges did Marie Curie face as a woman in science? Why is it important to balance discovery with safety? 4. Galileo Galilei: The Star Watcher Who Defied the Church Born: February 15, 1564 Died: January 8, 1642 Galileo loved looking at the stars. He built one of the first powerful telescopes and made stunning discoveries: mountains on the Moon, moons around Jupiter, and that the Earth orbits the Sun—not the other way around. This idea, called heliocentrism, went against the teachings of the Church. He was put on trial and forced to say he was wrong. But he wasn’t. He spent his last years under house arrest, quietly writing. Today, Galileo is called the father of modern science for daring to question what others blindly believed. Think About It: Why do you think Galileo was punished for telling the truth? Should science always follow evidence, even if it goes against powerful beliefs? 5. Isaac Newton: The Man Who Asked “Why?” When an Apple Fell Born: January 4, 1643 Died: March 31, 1727 One day, an apple fell from a tree, and Isaac Newton began to wonder: Why did it fall down, not sideways or up? This simple question led to his theory of gravity. Newton also invented calculus, described the laws of motion, and changed physics forever. But Newton wasn’t just a genius—he was curious, quiet, and often worked alone. He believed everything in nature followed rules, and it was our job to discover them. Thanks to him, we understand how planets move, how rockets launch, and why you fall when you trip. Think About It: How did Newton’s curiosity lead to great discoveries? Do you think working alone helped or hurt Newton? 6. Ada Lovelace: The First Computer Programmer Before Computers Existed Born: December 10, 1815 Died: November 27, 1852 Ada Lovelace was the daughter of the famous poet Lord Byron, but she didn’t love poetry—she loved numbers! At a time when girls were expected to sew, Ada studied mathematics. She met Charles Babbage, who designed an early computer called the Analytical Engine. Ada imagined the machine could do more than just math—it could create music, art, and even write! She wrote what is now considered the first computer program, long before real computers were built. Think About It: How did Ada imagine something that didn’t exist yet? Why do we call her a pioneer in technology? 7. Albert Einstein: The Man Who Brought Time and Space Together Born: March 14, 1879 Died: April 18, 1955 Albert Einstein wasn’t always a good student. In fact, his teachers thought he was slow. But Einstein thought deeply. He asked big questions like, “What if you could ride a beam of light?” His theories of relativity changed how we see space, time, and gravity. He also warned the world about the dangers of nuclear weapons, even though his ideas helped create them. Einstein believed science should help people, not harm them. With his messy hair, kind smile, and brilliant mind, he remains a symbol of genius. Think About It: Can someone be bad in school but still be brilliant? Should scientists be responsible for how their inventions are used? 8. Pythagoras: The Musician Who Loved Math Born: Around 570 BC Died: Around 495 BC Long ago in ancient Greece, Pythagoras believed the universe followed numbers. He discovered the Pythagorean Theorem, a rule about triangles that helps us build houses, design computers, and navigate space. He also believed that music had math inside it—that certain notes made perfect harmony because of mathematical ratios. Pythagoras started a secret school and taught his students to search for truth through numbers, shapes, and sound. Think About It: Why do you think Pythagoras saw math in everything? How does music relate to math? 9. Rosalind Franklin: The Woman Behind the DNA Discovery Born: July 25, 1920 Died: April 16, 1958 Rosalind Franklin loved looking closely at things. She used a special machine called X-ray crystallography to photograph molecules. One of her greatest photos, called Photo 51, showed the shape of DNA, the molecule that carries life’s instructions. But her work was taken without credit. Two men, Watson and Crick, used her photo to build their famous model of DNA and won the Nobel Prize. Rosalind died young and never knew how important her work became. Think About It: Why is it important to give credit in science? What can we learn from Rosalind’s quiet strength? 10. Carl Linnaeus: The Man Who Gave Names to Everything Born: May 23, 1707 Died: January 10, 1778 Have you ever wondered why a tiger is called Panthera tigris? That’s thanks to Carl Linnaeus, a Swedish scientist who created a way to name and organize every living thing. His system is still used today in biology. Linnaeus loved nature and spent his life collecting plants, animals, and even rocks. He believed that by organizing life, we could better understand it. Thanks to him, we now have a global “dictionary of nature.” Think About It: Why is it important to name and organize living things? How does order help us understand the world?
Southeast Asia, vast region of Asia situated east of the Indian subcontinent and south of China. It consists of two dissimilar portions: a continental projection (commonly called mainland Southeast Asia) and a string of archipelagoes to the south and east of the mainland (insular Southeast Asia). Extending some 700 miles (1,100 km) southward from the mainland into insular Southeast Asia is the Malay Peninsula; this peninsula structurally is part of the mainland, but it also shares many ecological and cultural affinities with the surrounding islands and thus functions as a bridge between the two regions. Mainland Southeast Asia is divided into the countries of Cambodia, Laos, Myanmar (Burma), Thailand, Vietnam, and the small city-state of Singapore at the southern tip of the Malay Peninsula; Cambodia, Laos, and Vietnam, which occupy the eastern portion of the mainland, often are collectively called the Indochinese Peninsula. Malaysia is both mainland and insular, with a western portion on the Malay Peninsula and an eastern part on the island of Borneo. Except for the small sultanate of Brunei (also on Borneo), the remainder of insular Southeast Asia consists of the archipelagic nations of Indonesia and the Philippines. Southeast Asia stretches some 4,000 miles at its greatest extent (roughly from northwest to southeast) and encompasses some 5,000,000 square miles (13,000,000 square km) of land and sea, of which about 1,736,000 square miles is land. Mount Hkakabo in northern Myanmar on the border with China, at 19,295 feet (5,881 meters), is the highest peak of mainland Southeast Asia. Although the modern nations of the region are sometimes thought of as being small, they are—with the exceptions of Singapore and Brunei—comparatively large. Indonesia, for example, is more than 3,000 miles from west to east (exceeding the west-east extent of the continental United States) and more than 1,000 miles from north to south; the area of Laos is only slightly smaller than that of the United Kingdom; and Myanmar is considerably larger than France. All of Southeast Asia falls within the tropical and subtropical climatic zones, and much of it receives considerable annual precipitation. It is subject to an extensive and regular monsoonal weather system (i.e., one in which the prevailing winds reverse direction every six months) that produces marked wet and dry periods in most of the region. Southeast Asia’s landscape is characterized by three intermingled physical elements: mountain ranges, plains and plateaus, and water in the form of both shallow seas and extensive drainage systems. Of these, the rivers probably have been of the greatest historical and cultural significance, for waterways have decisively shaped forms of settlement and agriculture, determined fundamental political and economic patterns, and helped define the nature of Southeast Asians’ worldview and distinctive cultural syncretism. It also has been of great importance that Southeast Asia, which is the most easily accessible tropical region in the world, lies strategically astride the sea passage between East Asia and the Middle Eastern–Mediterranean world. Within this broad outline, Southeast Asia is perhaps the most diverse region on Earth. The number of large and small ecological niches is more than matched by a staggering variety of economic, social, and cultural niches Southeast Asians have developed for themselves; hundreds of ethnic groups and languages have been identified. Under these circumstances, it often is difficult to keep in mind the region’s underlying unity, and it is understandable that Southeast Asia should so often be treated as a miscellaneous collection of cultures that simply do not quite fit anywhere else. Roofs of the Forbidden City, Beijing, China Britannica Quiz All About Asia Yet from ancient times Southeast Asia has been considered by its neighbors to be a region in its own right and not merely an extension of their own lands. The Chinese called it Nanyang and the Japanese Nan’yō, both names meaning “South Seas,” and South Asians used such terms as Suvarnabhūmi (Sanskrit: “Land of Gold”) to describe the area. Modern scholarship increasingly has yielded evidence of broad commonalities uniting the peoples of the region across time. Studies in historical linguistics, for example, have suggested that the vast majority of Southeast Asian languages—even many of those previously considered to have separate origins—either sprang from common roots or have been long and inseparably intertwined. Despite inevitable variation among societies, common views of gender, family structure, and social hierarchy and mobility may be discerned throughout mainland and insular Southeast Asia, and a broadly common commercial and cultural inheritance has continued to affect the entire region for several millennia. These and other commonalities have yet to produce a conscious or precise Southeast Asian identity, but they have given substance to the idea of Southeast Asia as a definable world region and have provided a framework for the comparative study of its components.
1.1945-1949: The immediate years after the Second World War ● At the end of 1945, Mao Zedong had come to see the USA as the greatest threat to his aspirations. a. He understood that East Asians were looking to the USA as the true liberator from Japanese imperialism. b. The USA’s support for the Kuomintang(KMT) and the restoration of U.S. authority in formerly Japanese Manchuria clashed with the CCP’s plans to use the region for its own needs in the impending civil war between the CCP and the GMD. ■ To compound matters, while the KMT was recognised internationally as the official government in China, Mao and the CCP saw the party as a puppet of U.S. imperialism. ● While Mao saw the USA as the greater threat to the CCP’s plans, Soviet actions also frustrated him. a. The USSR provided minimal and incoherent support for the Chinese Communists in Yan’an and Manchuria. b. Stalin also attempted to extract territorial and economic concessions from the Guomindang government in the Friendship and Alliance Treaty China signed in August 1945 under American and Soviet pressure in exchange for Soviet entry into the Second World War against Japan. ● The emerging superpower conflict over Europe and over American intervention in the impending civil war in China led to Mao’s ideological perception of the 8838/01 H1 History Paper 1 Theme II: The Cold War and East Asia (1945-1991) \ Page | 8 USA as an aggressive imperialist power that was hostile towards other countries, especially the USSR and China. ● In 1946, Mao promoted the theory of the intermediate zone, which envisioned a global united front against American imperialism. a. Mao saw the emerging superpower conflict as an American-Soviet contest for the intermediate zones, the capitalist, colonial and semi- colonial countries of West Europe, Africa, and Asia. b. Mao believed that the USSR was the defender of world peace. c. The intermediate zone, which included China, would not be part of the socialist camp. d. Despite the tremendous potential that U.S. aid held for China’s reconstruction, Mao’s ideological worldview and the impending civil war against the Guomindang prevented him from seeking normalised relations with the USA. In 1949, Mao decided to lean towards the side of the USSR despite two decades of unreliable support from them. e. Mao saw the anti-bourgeois campaigns in East Europe as evidence that China should isolate capitalist-bourgeois forces within it.2 f. Stalin had expelled Yugoslavia from the socialist camp as its leader, Tito was seen to have directly challenged Stalin’s authority. ■ Mao thus saw it as imperative to stress close unity to the USSR lest he was seen as a second Josip Broz Tito. At the same time, Mao sought a loose partnership with the USSR because Mao believed that China should preserve a high measure of self- reliance and zili gengsheng (自力更生) (regeneration through one’s own efforts). ● When the People’s Republic of China was formed on 1 October, 1949, relations between China’s and the USSR’s communists had improved substantially. a. However, the Chinese Communist Party (CCP) was also aware that the USSR never treated Chinese interests as a priority. What the CCP failed to fully understand was that Stalin ruled East Europe much like it was his empire and how this would have implications for China. b. In Mao’s first visit to the USSR in December 1949, Stalin was non- committal regarding the interests raised by the Chinese, and treated Mao as an underling as he feared that closer relations with the PRC would cause the USSR to lose privileges gained from the KMT. _________________________ 2 What Mao did not realise at that point was that the anti-bourgeois campaigns in East European countries were part of Stalin’s intentional design to consolidate the power of communists in them. 8838/01 H1 History Paper 1 Theme II: The Cold War and East Asia (1945-1991) \ Page | 9 A note on Sino-American relations 2. Early 1950: The USA’s hands-off policy towards Taiwan begins to change ● By early 1950, the Truman administration had written off Taiwan and believed it was only a matter of time before the island fell to the PLA. ● Two events in early 1950 changed the USA’s position on East Asia. ○ The formation of the USSR-PRC alliance in February 1950 ○ The North Korean invasion of South Korea in June 1950 3. 1950: The Sino-Soviet Friendship, Alliance and Mutual Assistance Treaty ● Signed on 14 February, 1950. 3.1Implications for Sino-Soviet relations ● Stalin saw it as a means to get concessions that he had failed to get from the Kuomintang (KMT) government in 1945. ● For Mao and the newly founded People’s Republic of China (PRC), the alliance would provide security against U.S. imperialism and allow the PRC to get economic aid for reconstruction from the USSR. ● The Chinese realised soon after the 1950 treaty had been signed that the Soviet Union was intent on exploiting the agreement in its own favour. 8838/01 H1 History Paper 1 Theme II: The Cold War and East Asia (1945-1991) \ Page | 10 ● The Sino-Soviet alliance was officially directed against Japanese militarism and its allies, especially the USA. ● The Sino-Soviet alliance comprised three elements: party, military and economic relations. ○ Party: The Chinese Communist Party (CCP) was included in the customs of communist party internationalism, such as regular exchange of party delegations to congresses of the fraternal parties in Stalin’s socialist camp. ■ This move was meant to bring the PRC’s ideological beliefs about communism into greater alignment with the USSR’s. ○ Military: The alliance was supposed to provide the newly formed and weak PRC with a strategic deterrent and military aid against the USA on three fronts: Guomindang-held Taiwan, divided Korea, and Vietnam where France attempted to reestablish its colonial control. ■ Convinced that the USA would aggressively seek ways to undermine the CCP-led PRC through Taiwan, Korea and Vietnam, Mao sought an active defence. ● While in Moscow, Mao unsuccessfully asked Stalin to provide military assistance for the liberation of Taiwan. ● At the beginning of 1950, the PRC delivered large-scale military aid to Hanoi. The PRC was the first country to grant the communist-led Democratic Republic of Vietnam diplomatic recognition on 18 January 1950; Mao persuaded Stalin to do so on 30 January 1950. ● The PRC committed itself to North Korea, where Mao saw the commitment to North Korea both as a defence against U.S. imperialism and as support for a fellow communist country. ○ Economic: During Mao’s first stay in Moscow, Stalin had personally promised the delivery of fifty projects for primary industrialisation. ■ The agreement also led to a series of supplementary ones, such as a US$ 300 million loan that the PRC would repay with a mixture of strategic materials, rubber, agricultural products, goods for daily use and hard currency. ■ Significantly, Stalin used Soviet military and economic aid to extract concessions similar to those he failed to get from the Guomindang government in 1945. ■ The USSR and PRC would disagree on the pace and extent of the PRC’s planned development. ● In the last five weeks of Stalin’s life in early 1953, he attempted to pressure the PRC to reduce the planned 8838/01 H1 History Paper 1 Theme II: The Cold War and East Asia (1945-1991) \ Page | 11 development speed to a mere annual growth of 13-14 percent, and to plan individual projects in detail beforehand. These moves would potentially result in the PRC’s economy growing at a slower rate than initially projected. ● However, after Stalin’s death on 5 March 1953, the PRC’s Zhou Enlai decided to use his visit of condolence to the USSR to press forward negotiations. ○ When talks resumed in 1 April 1953, Beijing pressed for 150 Soviet industrial projects, but Moscow reduced them to 91 on the basis of insufficient data provided by the Chinese. ■ The economic disarray after China’s civil war and the economic pressures that came with the Korean War influenced recovery and reconstruction in the early years of the PRC. ● Despite the PRC being unable to tap into Soviet economic assistance immediately, mutual trade between China and the USSR nevertheless increased 6.5 times from 1950 to 1956. ● Together with the 50 projects promised by Stalin in 1950, the final version of the First FYP for the PRC included 141 Soviet and 68 East European projects in a total of 649 planned. Three thousand Soviet advisers sent to China in subsequent years were directly linked to the First FYP. ● By 1955, over 60 percent of China’s goods exchange was with the USSR. ● Soviet economic assistance to China added up to the largest foreign development venture in the socialist camp ever. ○ The total number of planned projects amounted to between 300 and 360 projects. ○ However, the number of total finished projects ranged between 134 and 150. ● Transfers of knowledge and expertise were important to China’s economic development. ○ A study on Soviet experts counts 1,445 political advisers and 9,313 technical specialists sent to China until their sudden withdrawal in mid-1960. ■ For political reasons, the gradual withdrawal of advisers began after late 1956.
SS Spanish Version 8/25/25 Topic: Fall of Rome, Medieval Europe, and the Role of Monasteries Tema: La caída de Roma, la Europa medieval y el papel de los monasterios Reading Passage / Pasaje de Lectura The Roman Empire, once the most powerful civilization in the world, began to weaken during the 4th and 5th centuries. Several causes contributed to its decline, including political corruption, heavy taxes, reliance on slave labor, and invasions by barbarian tribes such as the Visigoths and Vandals. In 476 CE, the last Roman emperor in the West was removed, marking the official fall of the Western Roman Empire. This event brought significant changes to Europe, as centralized government disappeared and smaller kingdoms took control. El Imperio Romano, una vez la civilización más poderosa del mundo, comenzó a debilitarse durante los siglos IV y V. Varias causas contribuyeron a su declive, incluyendo la corrupción política, los altos impuestos, la dependencia de la mano de obra esclava y las invasiones de tribus bárbaras como los visigodos y los vándalos. En el año 476 d.C., el último emperador romano en Occidente fue depuesto, marcando la caída oficial del Imperio Romano Occidental. Este evento trajo cambios significativos a Europa, ya que el gobierno centralizado desapareció y reinos más pequeños tomaron el control. After the fall of Rome, Europe entered the Middle Ages, also known as Medieval Europe. This period lasted roughly from 500 to 1500 CE. Life during this time was shaped by the feudal system, where kings gave land to nobles in exchange for loyalty and military service. Most people were peasants who farmed the land and gave a portion of their harvest to their lords. Castles provided protection, while the Catholic Church became the most powerful institution, guiding people’s beliefs and daily lives. Después de la caída de Roma, Europa entró en la Edad Media, también conocida como la Europa medieval. Este período duró aproximadamente del año 500 al 1500 d.C. La vida durante este tiempo estaba organizada por el sistema feudal, en el cual los reyes daban tierras a los nobles a cambio de lealtad y servicio militar. La mayoría de las personas eran campesinos que cultivaban la tierra y entregaban una parte de su cosecha a sus señores. Los castillos brindaban protección, mientras que la Iglesia Católica se convirtió en la institución más poderosa, guiando las creencias y la vida diaria de las personas. In the uncertain times of Medieval Europe, monasteries served as centers of learning and stability. Monks lived simple lives dedicated to prayer, work, and study. They carefully copied ancient texts, preserving knowledge from Greece and Rome. Monasteries also offered medical care, shelter to travelers, and food to the poor. In many ways, they became islands of peace and knowledge in a world often filled with war and hardship. En los tiempos inciertos de la Europa medieval, los monasterios sirvieron como centros de aprendizaje y estabilidad. Los monjes vivían vidas simples dedicadas a la oración, el trabajo y el estudio. Ellos copiaban cuidadosamente textos antiguos, preservando el conocimiento de Grecia y Roma. Los monasterios también ofrecían atención médica, refugio a viajeros y comida a los pobres. De muchas maneras, se convirtieron en islas de paz y conocimiento en un mundo a menudo lleno de guerras y dificultades.
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