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Common Proteins and Genes in Cancer Biology
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Make a test, with answers best on the following: Conduct an investigation to provide evidence that living things are made of cells; either one cell or many different numbers and types of cells. Supporting Content LS1.A: Structure and Function • All living things are made up of cells, which is the smallest unit that can be said to be alive. An organism may consist of one single cell (unicellular) or many different numbers and types of cells (multicellular). (MS-LS-1.1) Further Explanation: Emphasis is on developing evidence that living things are made of cells, distinguishing between living and non-living things, and understanding that living things may be made of one cell or many and varied cells. In multicellular organisms, the body is a system of multiple interacting subsystems. These subsystems are groups of cells that work together to form tissues and organs that are specialized for particular body functions. (MS-LS-1.3) Further Explanation: Emphasis is on the conceptual understanding that cells form tissues and tissues form organs specialized for particular body functions. Examples could include the interaction of subsystems within a system and the normal functioning of those systems. Organisms reproduce, either sexually or asexually, and transfer their genetic information to their offspring. (MS-LS-1.4) • Living things share certain characteristics. (These include response to environment, reproduction, energy use, growth and development, life cycles, made of cells, etc.) (MS-LS1.4) Further Explanation: Examples should include both biotic and abiotic items, and should be defended using accepted characteristics of life. Plants, algae (including phytoplankton), and many microorganisms use the energy from light to make sugars (food) from carbon dioxide from the atmosphere and water through the process of photosynthesis, which also releases oxygen. These sugars can be used immediately or stored for growth or later use. (MS-LS-1.5) Further Explanation: Emphasis is on tracing movement of matter and flow of energy. Supporting Content LS1.C: Organization for Matter and Energy Flow in Organisms • Within individual organisms, food moves through a series of chemical reactions (cellular respiration) in which it is broken down and rearranged to form new molecules, to support growth, or to release energy. (MS-LS-1.6) Further Explanation: Emphasis is on describing that molecules are broken apart and put back together and that in this process, energy is released and on understanding that the elements in the products are the same as the elements in the reactants. Organisms, and populations of organisms, are dependent on their environmental interactions both with other living things and with nonliving factors. (MS-LS-2.1) • In any ecosystem, organisms and populations with similar requirements for food, water, oxygen, or other resources may compete with each other for limited resources, access to which consequently constrains their growth and reproduction. (MS-LS-2.1) • Growth of organisms and population increases are limited by access to resources. (MS-LS-2.1) Further Explanation: Emphasis is on cause and effect relationships between resources and growth of individual organisms and the numbers of organisms in ecosystems during periods of abundant and scarce resources. Similarly, predatory interactions may reduce the number of organisms or eliminate whole populations of organisms. Mutually beneficial interactions, in contrast, may become so interdependent that each organism requires the other for survival. Although the species involved in these competitive, predatory, and mutually beneficial interactions vary across ecosystems, the patterns of interactions of organisms with their environments, both living and nonliving, are shared. (MS-LS-2.2) Further Explanation: Emphasis is on predicting consistent patterns of interactions in different ecosystems in terms of the relationships among and between organisms and abiotic components of ecosystems. Examples of types of interactions could include competitive, predatory, and mutually beneficial. Food webs are models that demonstrate how matter and energy is transferred between producers, consumers, and decomposers as the three groups interact within an ecosystem. Transfers of matter into and out of the physical environment occur at every level. Decomposers recycle nutrients from dead plant or animal matter back to the soil in terrestrial environments or to the water in aquatic environments. The atoms that make up the organisms in an ecosystem are cycled repeatedly between the living and nonliving parts of the ecosystem. (MS-LS-2.3) Further Explanation: Emphasis is on describing the conservation of matter and flow of energy into and out of various ecosystems, and on defining the boundaries of the system. Ecosystems are dynamic in nature; their characteristics can vary over time. Disruptions to any physical or biological component of an ecosystem can lead to shifts in all its populations. (MSLS-2.5) Further Explanation: Emphasis is on recognizing patterns in data and making warranted inferences about changes in populations, and on evaluating empirical evidence supporting arguments about changes to ecosystems. Biodiversity describes the variety of species found in Earth’s terrestrial and oceanic ecosystems. The completeness or integrity of an ecosystem’s biodiversity is often used as a measure of its health. (MS-LS-2.6) Supporting Content LS4.D: Biodiversity • Changes in biodiversity can influence humans’ resources, such as food, energy, and medicines, as well as ecosystem services that humans rely on—for example, water purification and recycling. (MS-LS-2.6) Supporting Content ETS1.B: Developing Possible Solutions • There are systematic processes for evaluating solutions with respect to how well they meet the criteria and constraints of a problem. (MS-LS-2.6) Further Explanation: Examples of ecosystem services could include water purification, nutrient recycling, and prevention of soil erosion. Examples of design solution constraints could include scientific, economic, and social considerations. Genes are located in the chromosomes of cells, with each chromosome pair containing two variants of each of many distinct genes. Each distinct gene chiefly controls the production of specific proteins, which in turn affects the traits of the individual. Structural changes to genes (mutations) can result in changes to proteins, which can affect the structures and functions of the organism and thereby change traits. (MS-LS-3.1) Supporting Content LS3.B: Variation of Traits • In addition to variations that arise from sexual reproduction, genetic information can be altered because of mutations. Though rare, mutations may result in significant changes to the structure and function of proteins. Changes can be beneficial, harmful, or neutral to the organism. (MS-LS-3.1) Further Explanation: Emphasis is on conceptual understanding that changes in genetic material may result in making different proteins. Organisms reproduce, either sexually or asexually, and transfer their genetic information to their offspring. (MS-LS-3.2) Supporting Content LS3.A: Inheritance of Traits • Variations of inherited traits between parent and offspring arise from genetic differences that result from the subset of chromosomes (and therefore genes) inherited. (MS-LS-3.2) Supporting Content LS3.B: Variation of Traits • In sexually reproducing organisms, each parent contributes half of the genes acquired (at random) by the offspring. Individuals have two of each chromosome and hence two alleles of each gene, one acquired from each parent. These versions may be identical or may differ from each other. (MS-LS-3.2) Further Explanation: Emphasis is on using models such as simple Punnett squares and pedigrees, diagrams, and simulations to describe the cause and effect relationship of gene transmission from parent(s) to offspring and resulting genetic variation. The collection of fossils and their placement in chronological order is known as the fossil record and documents the change of many life forms throughout the history of the Earth. Anatomical similarities and differences between various organisms living today and between living and once living organisms in the fossil record enable the classification of living things. (MS-LS-4.1, MS-LS-4.2) Further Explanation: Emphasis is on finding patterns of changes in the level of complexity of anatomical structures in organisms and the chronological order of fossil appearance in the rock layers. The collection of fossils and their placement in chronological order is known as the fossil record and documents the change of many life forms throughout the history of the Earth. Anatomical similarities and differences between various organisms living today and between living and once living organisms in the fossil record enable the classification of living things. (MS-LS-4.1, MS-LS-4.2) Further Explanation: Emphasis is on explanations of the relationships among organisms in terms of similarity or differences of the gross appearance of anatomical structures. Scientific genus and species level names indicate a degree of relationship. (MS-LS-4.3) Further Explanation: Emphasis is on inferring general patterns of relatedness among structures of different organisms by comparing diagrams, pictures, specimens, or fossils. Natural selection leads to the predominance of certain traits in a population, and the suppression of others. (MS-LS-4.4) Further Explanation: Emphasis is on using concepts of natural selection, including overproduction of offspring, passage of time, variation in a population, selection of favorable traits, and heritability of traits. In artificial selection, humans have the capacity to influence certain characteristics of organisms by selective breeding. One can choose desired parental traits determined by genes, which are then passed to offspring. (MS-LS-4.5) Further Explanation: Emphasis is on identifying and communicating information from reliable sources about the influence of humans on genetic outcomes in artificial selection (such as genetic modification, animal husbandry, gene therapy), and on the influence these technologies have on society as well as the technologies leading to these scientific discoveries. Adaptation by natural selection acting over generations is one important process by which species change over time in response to changes in environmental conditions. Traits that support successful survival and reproduction in the new environment become more common; those that do not become less common. Thus, the distribution of traits in a population changes. (MS-LS-4.6) Further Explanation: Emphasis is on using mathematical models, probability statements, and proportional reasoning to support explanations of trends in changes to populations over time. Examples could include Peppered Moth population changes before and after the industrial revolution.
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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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. 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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 gover
Plant cells have three kinds of structures that are not found in animal cells and that are extremely important to plant survival: plastids, central vacuoles, and cell walls. PLANT CELLS Most of the organelles and other parts of the cell just described are common to all eukaryotic cells. However, plant cells have three additional kinds of structures that are extremely important to plant function: cell walls, large central vacuoles, and plastids. To understand why plant cells have structures not found in ani- mal cells, consider how a plant’s lifestyle differs from an animal’s. Plants make their own carbon-containing molecules directly from carbon taken in from the environment. Plant cells take carbon diox- ide gas from the air, and in a process called photosynthesis, they convert carbon dioxide and water into sugars. The organelles and structures in plant cells are shown in Figure 4-21. SECTION 4 OBJECTIVES ● List three structures that are present in plant cells but not in animal cells. ● Compare the plasma membrane, the primary cell wall, and the secondary cell wall. ● Explain the role of the central vacuole. ● Describe the roles of plastids in the life of a plant. ● Identify features that distinguish prokaryotes, eukaryotes, plant cells, and animal cells. VOCABULARY cell wall central vacuole plastid chloroplast thylakoid chlorophyll Chloroplast Golgi apparatus Mitochondrion Cell membrane Nucleolus Nucleus Cytoskeleton Rough endoplasmic reticulum Pore Smooth endoplasmic reticulum Central vacuole Ribosome Cell wall In addition to containing almost all of the types of organelles that animal cells contain, plant cells contain three unique features. Those features are the cell wall, the central vacuole, and plastids, such as chloroplasts. FIGURE 4-21 Copyright © by Holt, Rinehart and Winston. All rights reserved. 88 CHAPTER 4 CELL WALL The cell wall is a rigid layer that lies outside the cell’s plasma membrane. Plant cell walls contain a carbohydrate called cellulose. Cellulose is embedded in a matrix of proteins and other carbohy- drates that form a stiff box around each cell. Pores in the cell wall allow water, ions, and some molecules to enter and exit the cell. Primary and Secondary Cell Walls The main component of the cell wall, cellulose, is made directly on the surface of the plasma membrane by enzymes that travel along the membrane. These enzymes are guided by microtubules inside the plasma membrane. Growth of the primary cell wall occurs in one direction, based on the orientation of the microtubules. Other components of the cell wall are made in the ER. These materials move in vesicles to the Golgi and then to the cell surface. Some plants also produce a secondary cell wall. When the cell stops growing, it secretes the secondary cell wall between the plasma membrane and the primary cell wall. The secondary cell wall is very strong but can no longer expand. The wood in desks and tabletops is made of billions of secondary cell walls. The cells inside the walls have died and disintegrated. CENTRAL VACUOLE Plant cells may contain a reservoir that stores large amounts of water. The central vacuole is a large, fluid-filled organelle that stores not only water but also enzymes, metabolic wastes, and other materials. The central vacuole, shown in Figure 4-22, forms as other smaller vacuoles fuse together. Central vacuoles can make up 90 percent of the plant cell’s volume and can push all of the other organelles into a thin layer against the plasma membrane. When water is plentiful, it fills a plant’s vacuoles. The cells expand and the plant stands upright. In a dry period, the vacuoles lose water, the cells shrink, and the plant wilts. Other Vacuoles Some vacuoles store toxic materials. The vacuoles of acacia trees, for example, store poisons that provide a defense against plant-eating ani- mals. Tobacco plant cells store the toxin nicotine in a storage vacuole. Other vacuoles store plant pigments, such as the colorful pigments found in rose petals. The central vacuole occupies up to 90 percent of the volume of some plant cells. The central vacuole stores water and helps keep plant tissue firm. FIGURE 4-22 Central vacuole Nucleus Chloroplast Copyright © by Holt, Rinehart and Winston. All rights reserved. CELL STRUCTURE AND FUNCTION 89 PLASTIDS Plastids are another unique feature of plant cells. Plastids are organelles that, like mitochondria, are surrounded by a double mem- brane and contain their own DNA. There are several types of plastids, including chloroplasts, chromoplasts, and leucoplasts. Chloroplasts Chloroplasts use light energy to make carbohydrates from carbon dioxide and water. As Figure 4-23 shows, each chloroplast contains a system of flattened, membranous sacs called thylakoids. Thylakoids contain the green pigment chlorophyll, the main mole- cule that absorbs light and captures light energy for the cell. Chloroplasts can be found not only in plant cells but also in a wide variety of eukaryotic algae, such as seaweed. Chloroplast DNA is very similar to the DNA of certain photosyn- thetic bacteria. Plant cell chloroplasts can arise only by the divi- sion of preexisting chloroplasts. These facts may suggest that chloroplasts are descendants of ancient prokaryotic cells. Like mitochondria, chloroplasts are also thought to be the descendants of ancient prokaryotic cells that were incorporated into plant cells through a process called endosymbiosis. Chromoplasts Chromoplasts are plastids that contain colorful pigments and that may or may not take part in photosynthesis. Carrot root cells, for example, contain chromoplasts filled with the orange pigment carotene. Chromoplasts in flower petal cells contain red, purple, yellow, or white pigments. Other Plastids Several other types of plastids share the general features of chloro- plasts but differ in content. For example, amyloplasts store starch. Chloroplasts, chromoplasts, and amyloplasts arise from a common precursor, called a proplastid. Thylakoid Inner membrane Outer membrane chloroplast from the Greek chloros, meaning “pale green,” and plastos, meaning “formed” Word Roots and Origins A chloroplast captures energy from sunlight and uses that energy to convert carbon dioxide and water into sugar and other carbohydrates. FIGURE 4-23 Copyright © by Holt, Rinehart and Winston. All rights reserved. 90 CHAPTER 4 COMPARING CELLS All cells share common features, such as a cell membrane, cyto- plasm, ribosomes, and genetic material. But there is a high level of diversity among cells, as shown in Figure 4-24. There are signifi- cant differences between prokaryotes and eukaryotes. In addition, plant cells have features that are not found in animal cells. Prokaryotes Versus Eukaryotes Prokaryotes differ from eukaryotes in that prokaryotes lack a nucleus and membrane-bound organelles. Prokaryotes have a region, called a nucleoid, in which their genetic material is concen- trated. However, prokaryotes lack an internal membrane system. Plant Cells Versus Animal Cells Three unique features distinguish plant cells from animal cells. One is the production of a cell wall by plant cells. Plant cells contain a large central vacuole. Third, plant cells contain a variety of plastids, which are not found in animal cells. Cell walls, central vacuoles, and plastids are unique features that are important to plant function. 1. Identify three unique features of plant cells. 2. List the differences between the plasma mem- brane, the primary cell wall, and the secondary cell wall. 3. Identify three functions of plastids. 4. Name three things that may be stored in vacuoles. 5. Describe the features that distinguish prokary- otes from eukaryotes and plant cells from animal cells. CRITICAL THINKING
LESSON 2 Early Theories on the Origin of Life • Identify the different theories on the origin of life. • Describe each theory and determine their differences What are the characteristics of living things? Before learning about the history of Earth based on geological evidence, early scientists explored the possibilities of how the first life-form existed. There are several theories about the origin of life. Theory of Catastrophism The theory of catastrophism supported by French scientists Georges Cuvier (1769-1832) and Alcide Dessalines d'Orbigny (1802-1857), is said to be a modification of the creation story of the Bible. It states that there have been several living creations from God, each encountered a catastrophe that completely destroyed them. Each new creation consisted of new life-forms, which happen to be different from the previous ones. Theory of Abiogenesis The theory of abiogenesis, or the spontaneous generation theory, states that living things were naturally created from nonliving things such as simple organic compounds. The theory supposes that abiogenesis occurred between 3.8 and 4 Gya. The experiment performed by Stanley Miller in 1953 gave way to many speculations and studies on how life on Earth really began. His research involved a simulation of the possible environment on Earth in the past. He demonstrated how an electric spark (which is a simulation for lightning) when passed through simple organic gases (similar to the early Earth atmosphere), resulted in the formation of amino acids, which are now known as the building blocks of proteins and the components of living tissues. Theory of Biogenesis The theory of biogenesis presented a strong argument against abiogenesis. This theory states that living things come from living things. Experiments of Francesco Redi and Louis Pasteur disproved the thought of spontaneous generation during their time (figure 5-1). Louis Pasteur argued that life comes from preexisting life and not from nonliving material. However, it should be noted that the "abiogenesis" or "spontaneous generation" that he opposed referred to any modern, existing, fully-formed organisms, not the original generation of life. Louis Pasteur's law of biogenesis contradicted the common belief during his time that cattle dung gives rise to flies, or old clothes with rotten food gives rise to mice. The idea of spontaneous generation was popular until near the seventeenth century. Even famous scientists of that period, such as Descartes, Galileo, and Jan Baptista van Helmont, believed in this theory. CIENCE PIONEER Francesco Redi (1626-1697). Francesco Redi is a physician, a naturalist, and a poet. His works challenged the popular theory of spontaneous generation. He disproved the idea that living things may arise from nonliving things. He also worked on toxicology using viper's venom. He discovered and worked on some parasites that caused disease in humans. SCIENCE CAREER Evolutionary Biologist An evolutionary biologist studies the descent of species and the origin of new species of living things. Working as evolutionary biologist means studying and researching species diversity, their interaction with the environment, their adaptation to change, their ancestors, and their possible origins. This career is important in the field of science because it seeks an answer to the questions about how life began on Earth.
LARGE CARBON MOLECULES Many carbon compounds are built up from smaller, simpler molecules known as monomers (MAH-ne-mers), such as the ones shown in Figure 3-3. As you can also see in Figure 3-3, monomers can bond to one another to form polymers (PAWL-eh-mer). A polymer is a molecule that consists of repeated, linked units. The units may be identical or structurally related to each other. Large polymers are called macromolecules. There are many types of macromolecules, such as carbohydrates, lipids, proteins and nucleic acids. Monomers link to form polymers through a chemical reaction called a condensation reaction. Each time a monomer is added to a polymer, a water molecule is released. In the condensation reac- tion shown in Figure 3-4, two sugar molecules, glucose and fruc- tose, combine to form the sugar sucrose, which is common table sugar. The two sugar monomers become linked by a C—O—C bridge. In the formation of that bridge, the glucose molecule releases a hydrogen ion, H, and the fructose molecule releases a hydroxide ion, OH. The OH and H ions that are released then combine to produce a water molecule, H2O. In addition to building polymers through condensation reac- tions, living organisms also have to break them down. The break- down of some complex molecules, such as polymers, occurs through a process known as hydrolysis (hie-DRAHL-i-sis). In a hydrolysis reaction, water is used to break down a polymer. The water molecule breaks the bond linking each monomer. Hydrolysis is the reverse of a condensation reaction. The addition of water to some complex molecules, including polymers, under certain con- ditions can break the bonds that hold them together. For example, in Figure 3-4 reversing the reaction will result in sucrose breaking down into fructose and glucose. 2H2O Monomers Polymer C C O H OH C OH H CH2OH C H CH2OH C HO H C O H C OH H C CH2OH H C H OH O Sucrose C C O H OH C OH H CH2OH C H CH2OH C HO H C OH OH H C OH H C CH2OH H C H OH O Glucose Fructose H2O The condensation reaction below shows how glucose links with fructose to form sucrose. One water molecule is produced each time two monomers form a covalent bond. FIGURE 3-4 monomer from the Greek mono, meaning “single or alone,” and meros, meaning “a part” Word Roots and Origins A polymer is the result of bonding between monomers. In this example, each monomer is a six-sided carbon ring. The starch in potatoes is an example of a molecule that is a polymer. FIGURE 3-3 Copyright © by Holt, Rinehart and Winston. All rights reserved. 54 CHAPTER 3 ENERGY CURRENCY Life processes require a constant supply of energy. This energy is available to cells in the form of certain compounds that store a large amount of energy in their overall structure. One of these com- pounds is adenosine (uh-DEN-uh-SEEN) triphosphate, more commonly referred to by its abbreviation, ATP. The left side of Figure 3-5 shows a simplified ATP molecule struc- ture. The 5-carbon sugar, ribose, is represented by the blue carbon ring. The nitrogen-containing compound, adenine, is represented by the 2 orange rings. The three linked phosphate groups, —PO4 , are represented by the blue circles with a “P.” The phospate groups are attached to each other by covalent bonds. The covalent bonds between the phosphate groups are more unstable than the other bonds in the ATP molecule because the phosphate groups are close together and have negative charges. Thus, the negative charges make the bonds easier to break. When a bond between the phosphate groups is broken, energy is released. This hydrolysis of ATP is used by the cell to provide the energy needed to drive the chemical reactions that enable an organism to function.
CARBOHYDRATES Carbohydrates are organic compounds composed of carbon, hydrogen, and oxygen in a ratio of about one carbon atom to two hydrogen atoms to one oxygen atom. The number of carbon atoms in a carbohydrate varies. Some carbohydrates serve as a source of energy. Other carbohydrates are used as structural materials. Carbohydrates can exist as monosaccharides, disaccharides, or polysaccharides. Monosaccharides A monomer of a carbohydrate is called a monosaccharide (MAHN-oh-SAK-uh-RIED). A monosaccharide—or simple sugar— contains carbon, hydrogen, and oxygen in a ratio of 1:2:1. The gen- eral formula for a monosaccharide is written as (CH2O)n, where n is any whole number from 3 to 8. For example, a six-carbon mono- saccharide, (CH2O)6, would have the formula C6H12O6. The most common monosaccharides are glucose, fructose, and galactose, as shown in Figure 3-6. Glucose is a main source of energy for cells. Fructose is found in fruits and is the sweetest of the monosaccharides. Galactose is found in milk. Notice in Figure 3-6 that glucose, fructose, and galactose have the same molecular formula, C6H12O6, but differing structures. The different structures determine the slightly different properties of the three compounds. Compounds like these sugars, with a single chemical formula but different structural forms, are called isomers (IE-soh-muhrz). SECTION 2 OBJECTIVES ● Distinguish between monosaccharides, disaccharides, and polysaccharides. ● Explain the relationship between amino acids and protein structure. ● Describe the induced fit model of enzyme action. ● Compare the structure and function of each of the different types of lipids. ● Compare the nucleic acids DNA and RNA. VOCABULARY carbohydrate monosaccharide disaccharide polysaccharide protein amino acid peptide bond polypeptide enzyme substrate active site lipid fatty acid phospholipid wax steroid nucleic acid deoxyribonucleic acid (DNA) ribonucleic acid (RNA) nucleotide C HO H C H OH C OH H C CH2OH H C H OH O Glucose C OH C O H OH C OH H CH2OH C H CH2OH Fructose C H HO C OH H C OH H C CH2OH H C H OH O Galactose Glucose, fructose, and galactose have the same chemical formula, but their structural differences result in different properties among the three compounds. FIGURE 3-6 Copyright © by Holt, Rinehart and Winston. All rights reserved. 56 CHAPTER 3 Disaccharides and Polysaccharides In living things, two monosaccharides can combine in a condensa- tion reaction to form a double sugar, or disaccharide (die-SAK-e-RIED). For example in Figure 3-4, the monosaccharides fructose and glu- cose can combine to form the disaccharide sucrose. A polysaccharide is a complex molecule composed of three or more monosaccharides. Animals store glucose in the form of the polysaccharide glycogen. Glycogen consists of hundreds of glucose molecules strung together in a highly branched chain. Much of the glucose that comes from food is ultimately stored in your liver and muscles as glycogen and is ready to be used for quick energy. Plants store glucose molecules in the form of the polysaccha- ride starch. Starch molecules have two basic forms—highly branched chains that are similar to glycogen and long, coiled, unbranched chains. Plants also make a large polysaccharide called cellulose. Cellulose, which gives strength and rigidity to plant cells, makes up about 50 percent of wood. In a single cellu- lose molecule, thousands of glucose monomers are linked in long, straight chains. These chains tend to form hydrogen bonds with each other. The resulting structure is strong and can be broken down by hydrolysis only under certain conditions. PROTEINS Proteins are organic compounds composed mainly of carbon, hydrogen, oxygen, and nitrogen. Like most of the other biological macromolecules, proteins are formed from the linkage of monomers called amino acids. Hair and horns, as shown in Figure 3-7a, are made mostly of proteins, as are skin, muscles and many biological catalysts (enzymes). Amino Acids There are 20 different amino acids, and all share a basic structure. As Figure 3-7b shows, each amino acid contains a central carbon atom covalently bonded to four other atoms or functional groups. A single hydrogen atom, highlighted in blue in the illustration, bonds at one site. A carboxyl group, —COOH, highlighted in green, bonds at a second site. An amino group, —NH2, highlighted in yel- low, bonds at a third site. A side chain called the R group, high- lighted in red, bonds at the fourth site. The main difference among the different amino acids is in their R groups. The R group can be complex or it can be simple, such as the CH3 group shown in the amino acid alanine in Figure 3-7b. The differences among the amino acid R groups gives different proteins very different shapes. The different shapes allow pro- teins to carry out many different activities in living things. Amino acids are commonly shown in a simplified way such as balls, as shown in Figure 3-7c. (a) Many structures, such as hair and horns are made of proteins. (b) Proteins are made up of amino acids. Amino acids differ only in the type of R group (shown in red) they carry. Polar R groups can dissolve in water, but nonpolar R groups cannot. (c) Amino acids have complex structures, so, in this and other textbooks, they are often simplified into balls. FIGURE 3-7 (b) Alanine (an amino acid) (c) Simplified version of amino acid CH3 H N OH C C H O H (a) Copyright © by Holt, Rinehart and Winston. All rights reserved. BIOCHEMISTRY 57 H H N C C OH H O H CH3 H2O Glycine Alanine H N OH C C H O H H H N C C H O H CH3 N OH C C H O H (a) (b) (a) The peptide bond (shaded blue) that binds amino acids together to form a polypeptide results from a condensation reaction that produces water. (b) Poly- peptides are commonly shown as a string of balls in this textbook and elsewhere. Each ball represents an amino acid. FIGURE 3-8 Substrate Products Enzyme 1 2 3 In the induced fit model of enzyme action, the enzyme can attach only to a substrate (reactant) with a specific shape. The enzyme then changes and reduces the activation energy of the reaction so reactants can become products. The enzyme is unchanged and is available to be used again. 3 2 1 FIGURE 3-9 Dipeptides and Polypeptides Figure 3-8a shows how two amino acids bond to form a dipeptide (die-PEP-TIED). In this condensation reaction, the two amino acids form a covalent bond, called a peptide bond (shaded in blue in Figure 3-8a) and release a water molecule. Amino acids often form very long chains called polypeptides (PAHL-i-PEP-TIEDZ). Proteins are composed of one or more polypep- tides. Some proteins are very large molecules, containing hun- dreds of amino acids. Often, these long proteins are bent and folded upon themselves as a result of interactions—such as hydrogen bonding—between individual amino acids. Protein shape can also be influenced by conditions such as temperature and the type of solvent in which a protein is dissolved. For exam- ple, cooking an egg changes the shape of proteins in the egg white. The firm, opaque result is very different from the initial clear, runny material. Enzymes Enzymes—RNA or protein molecules that act as biological catalysts—are essential for the functioning of any cell. Many enzymes are proteins. Figure 3-9 shows an induced fit model of enzyme action. Enzyme reactions depend on a physical fit between the enzyme molecule and its specific substrate, the reactant being catalyzed. Notice that the enzyme has folds, or an active site, with a shape that allows the substrate to fit into the active site. An enzyme acts only on a specific substrate because only that substrate fits into its active site. The linkage of the enzyme and substrate causes a slight change in the enzyme’s shape. The change in the enzyme’s shape weakens some chemical bonds in the substrate, which is one way that enzymes reduce activation energy, the energy needed to start the reaction. After the reaction, the enzyme releases the products. Like any catalyst, the enzyme itself is unchanged, so it can be used many times. An enzyme may not work if its environment is changed. For example, change in temperature or pH can cause a change in the shape of the enzyme or the substrate. If such a change happens, the reaction that the enzyme would have catalyzed cannot occur.
• There are two groups of animals which are important in agriculture. • The groups are domestic and wild animals. 1. Domestic animals • These are animals which are trained to live with people in their homes. • Some examples includes : Uses of domestic animals • Source of food, for example milk, eggs and meat. • For cultural purposes, for example paying lobola. • Some are used for transport and labour. • Domestic animals can also be a source of income. 2 . Wild animals • These are animals which are found in game reserves and in the forests • They are also called game animals. • Examples of wild animals are: Uses of wild animals • Some of the wild animals give us meat, hides and ivory. • Wild animals attract visitors from other countries, so the country gets money. Wednesday 06 September 2023 Exercise: Domestic animals 1. What is a domestic animal? [2] 2. What is a wild animal? [2] 3. Name any 4 domestic animals that you know. [4] 4. State any 2 uses of domestic animals. [2] 5. Name any 3 wild animals that you know. [3] 6. Give 2 uses of wild ani mals. [2] 7. Wild animals can also be called animals. [1] • Animals, like human beings need good food to help them to grow and reproduce. • The main sources of food for animals include stock feeds, pastures, veld grass, crop remains and cereal grains. • These foods contain the most needed essential nutrients. Nutrient Function carbohydrates Gives energy Fats Give energy and warmth Proteins Helps in growth and repair of body parts Minerals Help in the formation of bones and teeth Vitamins Help develop good sight, improve fertility and help animals fight diseases Water Transports food in the body, cools the body and remove waste from the body. • Livestock are domesticated animals that are kept for food, use or for sale. • Small livestock has many uses. • They are a good source of food. • They also give us manure to use in gardens. • We get clothes and medicine as well from small livestock • Examples of small livestock are rabbits, sheep, goats and all form of poultry Poultry • All animals that are kept by farmers which have wings and feathers and lay eggs are called poultry. • They are a good source of white meat, eggs and manure. • They are also a source of income when we well them and their products. • Poultry includes chicken, guinea fowl, ducks, turkey, pigeons and quail birds. Rabbits • Rabbits have 3 known uses which are: They are kept for meat Kept for pelts. (pelt is animal skin used to make blankets, hats and jackets) They are used at agricultural shows • Rabbits are cheap to buy, easy to keep and feed. • They take about 4 months to mature. • Rabbits are fed using green vegetables and rabbit pellets. Wednesday 20 September 2023 Small livestock 1. Give 3 examples of small livestock. [3] 2. State 3 things that are provided by small livestock. [3] 3. What are the 2 uses of rabbits? [2] 4. Pelts are used to make _________________ [1] 5. State any 4 examples poultry. [4] 6. Rabbits take ___________ months to mature. [1] 7. What is poultry? • Apiculture is the keeping of bees in order for them to produce honey for sale. • Apiculture is very important because: (i) Provides honey - a valuable nutritional food (ii) Provides bees wax - which has many uses in industry Uses of wax For making candles Polish furniture Make crayons Prevent tools from rusting (iii) Honey bees are excellent pollinating agents, thus increasing agricultural yields. BEE COLONY Inhabitants of the bee colony and their roles • A honey bee colony typically consists of three kinds of adult bees: workers, drones and the queen 1. Workers • Workers are the smallest and constitute the majority of bees occupying the colony. • They do not lay eggs. • Workers have specialized structures, such as brood food glands, scent glands, wax glands, and pollen baskets. • these allow them to perform all the labors of the hive. Roles of the worker bees they forage for nectar, pollen, water, and plant sap. They clean and polish the cells. feed the brood. care for the queen. remove debris. handle incoming nectar. build beeswax combs. guard the entrance. 2. Drones • Drones (male bees) are the largest bees in the colony. • They are generally present only during late spring and summer. • The drone’s head is much larger than that of either the queen or worker. • Drones have no stinger, pollen baskets, or wax glands. • Their main function is to mate with the queen. 3 . Queen • Each colony has only one queen. • The queen is the largest of the bees in a bee colony. • The Queen Bee plays a vital role in the hive because she is the only female with fully developed ovaries. • She produces both fertilized and unfertilized eggs. • Queens lay the greatest number of eggs in the spring and early summer. • The queen also produce chemical scents that help regulate the unity of the colony. 1. What is apiculture? [2] 2. Give 3 reasons why apiculture is important in Zimbabwe. [3] 3. Name the 3 inhabitants of the bee colony. [3] 4. Briefly explain the roles of each inhabitant named in number 3. • Apart from using hand tools, farmers also use some farm implements and machinery to carry out their field work. • Machines help farmers do their work more easily and quickly. • The most common implements used by farmers to grow, harvest and transport their produces are: mouldboard plough Cultivator Scotch cart Harrow Planter Maize sheller combine harvesters Boom sprayers Disc harrow Spike toothed harrow KNAPSACK SPRAYER • Farming is a business. • Communal farmers grow crops and keep animal mainly for their own use. • If there is any extra they sell to get money. • Commercial farmers grow crops and keep animals for sale. • Crops and animals produced are called farm produce. • There are places were farmers have to sell their produce. • Farmers can take their produce to local markets. • A market is a place where buying and selling occurs. • Some of the local markets includes: A shopping centre A school A nearest bus stop A local village A school Local Grain Marketing Board depot (GMB) Types of farm produce Beef Fruits Mutton Eggs Vegetables Pork Milk Chicken Cereals/grains beans
Lipids are large, nonpolar organic molecules. They do not dissolve in water. Lipids include triglycerides (trie-GLIS-uhr-IEDZ), phospho- lipids, steroids, waxes, and pigments. Lipid molecules have a higher ratio of carbon and hydrogen atoms to oxygen atoms than carbohydrates have. Because lipid molecules have larger numbers of carbon-hydrogen bonds per gram than other organic com- pounds do, they store more energy per gram. Fatty Acids Fatty acids are unbranched carbon chains that make up most lipids. Figure 3-10 shows that a fatty acid contains a long carbon chain (from 12 to 28 carbons) with a carboxyl group, —COOH, attached at one end. The two ends of the fatty-acid molecule have different properties. The carboxyl end is polar and is thus hydrophilic or attracted to water molecules. In contrast, the hydro- carbon end of the fatty-acid molecule is nonpolar. This end tends not to interact with water molecules and is said to be hydrophobic (HIE-droh-FOH-bik), or “water fearing.” In saturated fatty acids, such as palmitic acid, which is shown in Figure 3-10, each carbon atom is covalently bonded to four atoms. The carbon atoms are in effect full, or saturated. In contrast, linoleic acid, also shown in Figure 3-10, has carbon atoms that are not bonded to the maximum number of atoms to which they can bond. Instead, they have formed double bonds within the carbon chain. This type of fatty acid is said to be unsaturated. Triglycerides Three classes of lipids important to living things contain fatty acids: triglycerides (fats), phospholipids, and waxes. A triglyceride is composed of three molecules of fatty acid joined to one molecule of the alcohol glycerol. Saturated triglycerides are composed of saturated fatty acids. They typically have high melting points and tend to be hard at room temperature. Common dietary saturated triglycerides include butter and fats in red meat. In contrast, unsaturated triglycerides are composed of unsaturated fatty acids and are usually soft or liquid at room temperature. Unsaturated triglycerides are found primarily in plant seeds where they serve as an energy and carbon source for germinating plants. Phospholipids Phospholipids have two, rather than three, fatty acids attached to a molecule of glycerol. They have a phosphate group attached to the third carbon of the glycerol. As shown in Figure 3-11, the cell membrane is made of two layers of phospholipids, called the lipid bilayer. The inability of lipids to dissolve in water allows the mem- brane to form a barrier between the inside and outside of the cell. Hydrophilic “head” Phospholipids Hydrophobic “tail” Phospholipids Water Water The lipid bilayer of a cell membrane is a double row of phospholipids.The “tails” face each other.The “head” of a phospholipid, which contains a phosphate group, is polar and hydrophilic.The two tails are two fatty acids and are nonpolar and hydrophobic. FIGURE 3-11 H C H C O OH H C H H C H H C H H C H H C H H C H H C H H C H H C H H C H H C H H C H C H H H H C H H C H H C H H C H H C H C O OH H C H H C H H C H C H C H C H H C H H C H C H H C H H C H C H H H H C H Fatty acids have a polar carboxyl head, highlighted in purple, and a nonpolar hydrocarbon tail, highlighted in green. FIGURE 3-10 Palmitic acid Linoleic acid mb06se_bchs02.qxd 5/18/07 10:49 AM Page 59 60 CHAPTER 3 1. Compare the structure of monosaccharides, dis- accharides, and polysaccharides. 2. How are proteins constructed from amino acids? 3. How do amino acids differ from one another? 4. Describe a model of enzyme action. 5. Why do phospholipids orient in a bilayer when in a watery environment, such as a cell? 6. Describe how the three major types of lipids differ in structure from one another. 7. What are the functions of the two types of nucleic acids? CRITICAL THINKING 8. Applying Information Before a long race, run- ners often “carbo load.” This means that they eat substantial quantities of carbohydrates. How might this help their performance? 9. Recognizing Relationships High temperatures can weaken bonds within a protein molecule. How might this explain the effects of using a hot curling iron or rollers in one’s hair? 10. Applying Information You want to eat more unsaturated than saturated fats. Name examples of foods you would eat more of and less of. SECTION 2 REVIEW Waxes A wax is a type of structural lipid consisting of a long fatty-acid chain joined to a long alcohol chain. Waxes are waterproof, and in plants, form a protective coating on the outer surfaces. Waxes also form protective layers in animals. For example, earwax helps pre- vent microorganisms from entering the ear canal. Steroids Unlike most other lipids, which are composed of fatty acids, steroid molecules are composed of four fused carbon rings with various functional groups attached to them. Many animal hor- mones, such as the male hormone testosterone, are steroid com- pounds. One of the most familiar steroids in humans is cholesterol. Cholesterol is needed by the body for nerve and other cells to func- tion normally. It is also a component of the cell membrane. NUCLEIC ACIDS Nucleic acids are very large and complex organic molecules that store and transfer important information in the cell. There are two major types of nucleic acids: deoxyribonucleic acid and ribonucleic acid. Deoxyribonucleic acid, or DNA, contains information that deter- mines the characteristics of an organism and directs its cell activi- ties. Ribonucleic (RIE-boh-noo-KLEE-ik) acid, or RNA, stores and transfers information from DNA that is essential for the manufactur- ing of proteins. Some RNA molecules can also act as enzymes. Both DNA and RNA are polymers, composed of thousands of linked monomers called nucleotides (NOO-klee-uh-TIEDS). As shown in Figure 3- 12, each nucleotide is made of three main components: a phosphate group, a five-carbon sugar, and a ring-shaped nitrogenous base.