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Title (Slide 0): "Digging Deeper: The Truth About Tillage" Subtitle: How turning the soil affects plants, microbes, and the planet Slide 1: What Is Tillage? Tilling the soil means digging, turning, and loosening it using tools or machines. It's a common farming practice to prepare the land before planting. Slide 2: Why Do Farmers Till? Tillage is usually done before planting to: • Soften and aerate the soil • Mix in nutrients • Remove weeds • Bury crop residues for decomposition and fertility Slide 3: Tools Used for Tillage Farmers use tools like: • Ploughs: Cut deep into the soil • Harrows: Break up clumps and smooth the surface Slide 4: Ploughs vs. Harrows • Ploughs: Used first, go deep, lift and flip soil • Harrows: Used after ploughs, work on the surface to break clumps and level the soil Slide 5: Types of Tillage Systems From most to least soil disturbance: • Conventional Tillage: Deep ploughing • Minimum Tillage: Light disturbance • Conservation Tillage: Only disturb seed zone, keep residues on top • Zero Tillage (No-Till): Plant directly into undisturbed soil Slide 6: Problem 1 – Soil Erosion Tillage removes protective cover, exposing soil to wind and rain. Result: topsoil—the most fertile layer—is easily washed or blown away. Slide 7: Problem 2 – Disruption of Soil Life Soil is a living ecosystem! • Worms, fungi, and bacteria help aerate soil and release nutrients • Tillage destroys their habitat, reducing fertility and soil health Slide 8: Problem 3 – Loss of Soil Structure Healthy soil has pores for air, water, and roots. Tillage breaks the sponge-like structure, and soil compacts over time—like flattening it into a pancake. Hard soil = poor plant growth. Slide 9: Problem 4 – Decreased Organic Matter Microbes "eat" organic matter through aerobic respiration (using O₂ and releasing CO₂). Tillage adds oxygen, microbes speed up, and burn through the soil’s “pantry” of organic matter—leaving it empty and poor. Slide 10: Problem 5 – Greenhouse Gas Emissions Faster decomposition = more CO₂ released. Tillage boosts microbial activity, which increases carbon dioxide emissions—contributing to climate change. ✅ Conclusion (Slide 11): 🌱 Tillage: A Double-Edged Tool Tillage can help prepare the soil and control weeds—but it comes at a cost. Over time, repeated tilling can strip away organic matter, destroy soil life, and release greenhouse gases. It's like spending all your savings for quick results—and being left with nothing for the future. The smarter path? Use reduced or no-till methods that protect soil health, keep carbon in the ground, and support long-term farming success.Key Word Definition
Aid Assistance in the form of grants or loans at below market rates. For example, the UK provided aid to Nepal after the 2015 earthquake to help with reconstruction efforts.
Barriers to Trade Government constraints on the flow of international goods and services, such as tariffs and quotas. For example, the European Union imposes tariffs on certain agricultural products to protect its farmers.
BRICS An acronym for an association of five major emerging nations: Brazil, Russia, India, China, and South Africa. For example, BRICS nations often meet to discuss economic cooperation and development strategies.
Demographic Aging The rise in the median age of a population. For example, Japan is experiencing demographic aging, with a significant increase in the elderly population.
Economic Core Region The most highly developed region(s) in a country. For example, London is considered an economic core region in the UK due to its high level of development and economic activity.
GNP (Gross National Product) GDP plus overseas earnings, also known as GNI (Gross National Income). For example, the GNP of the United States includes the value of goods and services produced domestically and the income earned by its citizens abroad.
HIC (Higher Income Country) A country with a high level of income and development. For example, Germany is classified as a higher income country due to its high GDP per capita and advanced infrastructure.
NEE (Newly Emerging Economy) A country that is experiencing rapid economic growth and industrialisation. For example, China is an example of a newly emerging economy, having rapidly industrialised and grown economically over the past few decades.
LIC (Lower Income Country) A country with a low level of income and development. For example, Malawi is considered a lower income country, with a low GDP per capita and limited access to healthcare and education.
GDHI (Gross Disposable Household Income) The amount of money that households have available for spending and saving after taxes and social contributions. For example, in the UK, the GDHI varies significantly between regions, with London having one of the highest levels.
Gini Coefficient A measure of income inequality within a population, ranging from 0 (perfect equality) to 100 (perfect inequality). For example, South Africa has a high Gini coefficient, indicating significant income inequality within the country.
Trade Unions Organisations that promote trade between member countries, such as the East African Community (EAC). For example, the East African Community (EAC) works to promote economic integration and trade among its member states.
Fair Trade A movement aimed at helping producers in developing countries achieve better trading conditions and promote sustainability. For example, Fairtrade coffee ensures that farmers receive a fair price for their product and work under safe conditions.
FDI (Foreign Direct Investment) Investment made by a company or individual in one country in business interests in another country. For example, Toyota's investment in manufacturing plants in the UK is an example of foreign direct investment.
Debt Relief The partial or total remission of debts, especially those owed by developing countries to external creditors. For example, the Heavily Indebted Poor Countries (HIPC) initiative provides debt relief to eligible countries to help them achieve sustainable development.
International Aid Voluntary transfer of resources from one country to another, often in the form of financial assistance, goods, or services. For example, the UK provides international aid to various countries through its Department for International Development (DFID).
Top-Down Development Large-scale development projects led by national governments or international organisations. For example, the construction of the Three Gorges Dam in China is an example of a top-down development project.
Bottom-Up Development Small-scale development projects led by local communities or NGOs, focusing on the needs of the poorest and most vulnerable. For example, WaterAid's installation of hand pumps in rural villages in Africa is an example of a bottom-up development project.
Urbanisation The increase in the proportion of people living in urban areas compared to rural areas. For example, rapid urbanisation in India has led to the growth of megacities like Mumbai and Delhi.
Geopolitics The study of the effects of geography (human and physical) on international politics and relations. For example, the geopolitics of the Arctic region involves disputes over territorial claims and access to natural resources.
Quality of Life The general well-being of individuals and societies, outlining negative and positive features of life. For example, Scandinavian countries are often ranked high in quality of life due to their strong social welfare systems and high levels of happiness.
Poverty Cycle A set of factors or events by which poverty, once started, is likely to continue unless there is outside intervention. For example, lack of education and healthcare can trap families in a poverty cycle, making it difficult for future generations to improve their living standards.
Sustainable Development Economic development that is conducted without depletion of natural resources. For example, the use of renewable energy sources like wind and solar power is a key aspect of sustainable development.
Humanitarian Aid Material or logistical assistance provided for humanitarian purposes, typically in response to crises including natural disasters and man-made disaster. For example, humanitarian aid was provided to Haiti after the devastating earthquake in 2010 to help with immediate relief efforts.
Economic Growth An increase in the production of goods and services in an economy over a period of time. For example, China's economic growth over the past few decades has lifted millions of people out of poverty.
Social Indicators Measures that describe the well-being of individuals or communities, such as health, education, and income. For example, life expectancy and literacy rates are common social indicators used to assess development.
Environmental Sustainability Responsible interaction with the environment to avoid depletion or degradation of natural resources and allow for long-term environmental quality. For example, practices like recycling and conservation of natural habitats contribute to environmental sustainability.
Infrastructure The basic physical and organisational structures and facilities needed for the operation of a society or enterprise. For example, good infrastructure, such as roads, bridges, and schools, is essential for economic development.
Globalisation The process by which businesses or other organisations develop international influence or start operating on an international scale. For example, the globalisation of technology companies like Apple and Google has led to their products being available worldwide.
Microfinance Financial services provided to low-income individuals or groups who are typically excluded from traditional banking. For example, microfinance institutions like Grameen Bank provide small loans to entrepreneurs in developing countries to help them start or expand their businesses.
Soils Southeast Asia, on balance, has a higher proportion of relatively fertile soils than most tropical regions, and soil erosion is less severe than elsewhere. Much of the region, however, is covered by tropical soils that generally are quite poor in nutrients. Often the profusion of plant life is more related to heat and moisture than to soil quality, even though these climatic conditions intensify both chemical weathering and the rate of bacterial action that usually improve soil fertility. Once the vegetation cover is removed, the supply of humus quickly disappears. In addition, the often heavy rainfall leaches the soils of their soluble nutrients, hastens erosion, and damages the soil texture. The leaching process in part results in laterites of reddish clay that contain hydroxides of iron and alumina. Laterite soils are common in parts of Myanmar, Thailand, and Vietnam and also occur in the islands of the Sunda Shelf, notably Borneo. The most fertile soils occur in regions of volcanic activity, where the ejecta is chemically alkaline or neutral. Such soils are found in parts of Sumatra and much of Java in Indonesia. The alluvial soils of the river valleys also are highly fertile and are intensively cultivated. Climate All of Southeast Asia falls within the warm, humid tropics, and its climate generally can be characterized as monsoonal (i.e., marked by wet and dry periods). Changing seasons are more associated with rainfall than with temperature variations. There is, however, a high degree of climatic complexity within the region. Temperatures Regional temperatures at or near sea level remain fairly constant throughout the year, although monthly averages tend to vary more with increasing latitude. Thus, with the exception of northern Vietnam, annual average temperatures are close to 80 °F (27 °C). Increasing elevation acts to decrease average temperatures, and such locations as the Cameron Highlands in peninsular Malaysia and Baguio in the Philippines have become popular tourist destinations in part because of their relatively cooler climates. Proximity to the sea also tends to moderate temperatures. Precipitation Much of Southeast Asia receives more than 60 inches (1,500 millimeters) of rainfall annually, and many areas commonly receive double and even triple that amount. The rainfall pattern is distinctly affected by two prevailing air currents: the northeast (or dry) monsoon and the southwest (or wet) monsoon. The northeast monsoon occurs roughly from November to March and brings relatively dry, cool air and little precipitation to the mainland. As the southwestward-flowing air passes over the warmer sea, it gradually warms and gathers moisture. Precipitation is especially heavy where the airstream is forced to rise over mountains or encounters a landmass. The east coast of peninsular Malaysia, the Philippines, and parts of eastern Indonesia receive the heaviest rains during this period. The southwest monsoon prevails from May to September, when the air current reverses and the dominant flow is to the northeast. The mainland receives the bulk of its rainfall during this period. Over much of the southern Malay Peninsula and insular Southeast Asia there is little or no prolonged dry season. This is especially marked in much of the equatorial region and along the east coast of the Philippines. While the dry and wet monsoons are important in explaining rainfall patterns, so too are such factors as relief, land and sea breezes, convectional overturning and cyclonic disturbances. These factors often are combined with monsoonal effects to produce highly variable rainfall patterns over relatively short distances. While many of the cyclonic disturbances produce only moderate rainfall, others mature into tropical storms—called cyclones in the Indian Ocean and typhoons in the Pacific—that bring heavy rains and destruction to the areas over which they pass. The Philippines are particularly affected by these storms. Plant life Tropical forests in Southeast Asia Tropical forests in Southeast Asia The seasonal nature and pattern of Southeast Asia’s rainfall, as well as the region’s physiography, have strongly affected the development of natural vegetation. The hot, humid climate and enormous variety of habitats have given rise to an abundance and diversity of vegetative forms unlike that in any other area of the world. Much of the natural vegetation has been modified by human action, although large areas of relatively untouched land still can be found. The vegetation can be grouped into two broad categories: the tropical-evergreen forests of the equatorial lowlands and the open type of tropical-deciduous, or “monsoon,” forests in areas of seasonal drought. The evergreen forests are characterized by multiple stories of vegetation, consisting of a variety of trees and plants. Although a large diversity of tree species is found in these forests, members of the Dipterocarpaceae family account for roughly half of the varieties. Deciduous forests are found in eastern Indonesia and those parts of the mainland where annual rainfall does not exceed 80 inches. Just as in the equatorial forest, a wide variety of species is normally the rule. Certain species, such as teak, have become highly valued commercially. Teak is found in parts of Indonesia, Myanmar, Thailand, and Laos. In addition to these two basic types of vegetation, other regional patterns reflect topography. Especially noteworthy are coastal and highland plant communities. Mangrove belts, of which there are more than 30 varieties, occur where silt is deposited in coastal areas. Upland forests dominated by maples, oaks, and magnolias are found especially on mainland mountain slopes. Human activity has been rapidly altering the stands of virgin forest in Southeast Asia. Most deforestation results from removal for fuelwood and clearing for agriculture and grazing. Although only a relatively small portion of the total land area has been permanently cleared for cultivation—e.g., in Java (Indonesia) and western Luzon (the Philippines)—in some areas shifting cultivation has brought about the replacement of virgin forest with secondary growth. In addition, nearly all countries have commercial logging industries; notable are those in Indonesia, Malaysia, Thailand, and Myanmar. A growing problem has been illegal logging. Thus, timber harvesting has come to contribute significantly to deforestation. Programs in social forestry and reforestation have yet to halt the rapid denuding of the landscape. Animal life Southeast Asia is situated where two major divisions of the world’s fauna meet. The region itself constitutes the eastern half of what is called the Oriental, or Indian, zoogeographic region (part of the much larger realm of Megagaea). Bordering along the south and east is the Australian zoogeographic region, and the eastern portion of insular Southeast Asia—Celebes (Sulawesi), the Moluccas, and the Lesser Sunda Islands—constitutes a transition zone between these two faunal regions. a classroom in Brazil More From Britannica education: Southeast Asia Southeast Asia is notable, therefore, for a considerable diversity of wildlife throughout the region. These differences are especially striking between the species of the eastern and western fringes as well as between those of the archipelagic south and the mainland north. The differences stem largely from the isolation, over varying lengths of geologic time, of species following their migration from the Asian continent. In addition, the tropical rain forests in many parts of the region, with their great diversity of vegetation, have made possible the development of complex communities of animals that fill specialized ecological niches. Especially numerous are arboreal and flying creatures. orangutans orangutansOrangutans (Pongo pygmaeus) in Sumatra, Indonesia. The distinction between the two faunal regions is best depicted by their mammal populations. In general, Australia is inhabited largely by marsupials (pouched mammals) and monotremes (egg-laying mammals), while Southeast Asia contains placental mammals and such hybrid species as the bandicoot of eastern Indonesia. Small mammals such as monkeys and shrews are the most numerous, while in many areas the larger mammals have been pushed into more remote areas and national preserves. Bears, gibbons, elephants, deer, civets, and pigs are found in both mainland and insular Southeast Asia, as are diminishing numbers of tigers. The Malayan tapir, a relative of the rhinoceros, is native to the Malay Peninsula and Sumatra, while the tarsier is found in the Philippines and parts of Indonesia. A number of rare endemic species are found in Indonesia and East (insular) Malaysia, including the Sumatran and Javan rhinoceros, the orangutan, the anoa (a dwarf buffalo), the babirusa (a wild swine), and the palm civet. As the pace of development accelerates and populations continue to expand in Southeast Asia, concern has increased regarding the impact of human activity on the region’s environment. A significant portion of Southeast Asia, however, has not changed greatly and remains an unaltered home to wildlife. The nations of the region, with only few exceptions, have become aware of the need to maintain forest cover not only to prevent soil erosion but to preserve the diversity of flora and fauna. Indonesia, for example, has created an extensive system of national parks and preserves for this purpose. Even so, such species as the Javan rhinoceros face extinction, with only a handful of the animals remaining in western JavaAccording to 〜によれば add to 増やす add up 合計する after a while しばらくして against the idea その考えに反対して all of a sudden 突然に all the time いつも all through the night 一晩中 along with 〜と一緒に apply for 〜に申し込む apply to 〜に適用する as a rule 原則として aside from 〜のほかに / 〜を除いて at any cost どんな犠牲を払っても at heart 心の底では at last ついに at least 少なくとも at length 詳細に at most 多くても at once すぐに / 一度に at the sight of 〜を見て at times 時々 attach to 〜に付ける / 〜に結びつける back up 支援する、バックアップする based on 〜に基づいて be absent from 〜を欠席している be against 反対する be based on 〜に基づいている be confident of 〜に自信がある be curious about 〜に好奇心を持つ be derived from 〜に由来する be filled with 〜で満たされている be full of 〜でいっぱいである be made up of 〜で構成されている be pleased with 〜に満足している be short of 不足している be similar to 〜に似ている because of 〜のせいで / 〜のおかげで before long まもなく break out 突発する break out in （急に）〜になる break up 解散する / 別れる bring out 引き出す / 公表する bring up 育てる / 持ち出す burst into 急に〜し始める by heart 暗記して by mistake 間違えて by now 今頃までには by the way ところで by way of 〜を通じて call for 要求する / 呼びかける call out 大声で呼ぶ carry on 続ける carry out 実行する、行う catch up with 〜に追いつく close to 〜に近い come across 偶然出会う / 見つける come into 〜に入る / 〜になる come out 出てくる / 公表される come up with 〜を思いつく compared with 〜と比べて depend on 〜に依存する do him good 彼に利益をもたらす drive at 意図する、狙う drop by 立ち寄る drop down 落ちる feel at home くつろぐ feel like doing 〜したい気分 feel sorry 気の毒に思う figure out 理解する find fault with 〜に文句をつける find out 知る、解明する for fear of 〜を恐れて for free 無料で for good 永遠に、完全に for once 一度だけ / 今回だけは for sale 売り物の for the best 最善のために get over 乗り越える get ready 準備する get rid of 〜を取り除く give away 与える、寄付する give in to 〜に屈する give off 放つ give out 配る / 発表する go ahead 続けて行う go down 下がる / 沈む hand in 提出する hand over 手渡す hang on 待つ / 頑張る hang up 受話器を置く hear from 〜から連絡をもらう help yourself 自由に取る / 自由にどうぞ hold back 控える hold on 待つ / 持ちこたえる hold up 停止させる、遅らせる in a jacket ジャケットを着て in a word 一言で言えば in advance 前もって in case 〜の場合に in common 共通して in detail 詳細に in hand 手元に in part 部分的に in place of 〜の代わりに in return 見返りに in terms of 〜の観点から in the distance 遠くに in the habit of 〜する習慣がある in the way 妨げになって instead of 〜の代わりに keep a secret 秘密を守る keep an eye on 見守る keep away from 近づかないようにする keep on Ving 〜し続ける keep pace with 〜に遅れずについていく keep the change お釣りはいりません keep track of 記録をつける / 追跡する keep up with 〜に遅れずについていく lay it down それを置く / 規定する less than 〜未満 / 〜より少ない look after 世話をする look back on 〜を振り返る look down on 〜を見下す look like 〜のように見える look up 調べる / 見上げる made up of 〜で構成されている major in 〜を専攻する make efforts 努力する make it out 理解する / 成し遂げる make out 見分ける、うまくいく make progress 進歩する make sense 理解できる、意味を成す make up your mind 決心する mind your own business 自分のことに集中しろ move on 次に進む no longer もはや〜ない not always 必ずしも〜でない nothing but ただ〜だけ on air 放送中 on behalf of 〜を代表して on business 仕事で on fire 火がついている on purpose わざと on the point 〜の点で on time 時間通りに one another お互いに out of the question 問題外で pass by 通り過ぎる pay attention 注意を払う play a part in 〜で役割を果たす pour out 注ぎ出す、溢れ出る prefer A to B BよりAを好む put away 片付ける put off 延期する put on 着る / 演じる (weightで太る） put out 消す / 発表する reach for 手を伸ばす rely on 〜に頼る result in 【自動詞】結果として〜になる run it over それをひく / 読み返す run out of 〜を使い果たす run over ひいてしまう、走り回る see about 手配する / 調べる see off 見送る see through 見抜く / 見通す set out 出発する、始める set up 設置する / 設定する show off 自慢する、見せびらかす show up 現れる / 到着する sit up 座る、起き上がる speak up はっきり話す stand out 目立つ stand out 目立つ stand up for 〜を支持する suffer from 〜に苦しむ take away 持ち去る / 奪う take in 理解する、取り入れる take notice of 注意を払う take on 引き受ける / 挑む take out 取り出す / 持ち帰る take over 引き継ぐ、乗っ取る take part 参加する take place 起こる / 開催される take risks 危険を冒す take turns 交代で行う talk over 相談する tear off 引き裂く the second largest 二番目に大きい think better of 考え直す throw away 捨てる try on 試着する turn in 提出する turn off 消す / 切る turn on （スイッチを）入れる turn out 結果的に〜になる turn over ひっくり返す under control 制御下にある up to date 最新の with ease 容易に with regard to 〜に関して 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. 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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. 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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 goverThe most common mistake made by Recruiter How well do you know the most common past participles in English?Past simple and past participle of the most common irregular verbs