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Secondary growth pt.3
Quiz by Morris Agata
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â These three layers -- phelloderm, cork cambium, and cork -- are referred to as a ??
âdie and are shed as secondary growth proceeds.
 These three layers -- phelloderm, cork cambium, and cork -- are referred to as a ??
die and are shed as secondary growth proceeds.
is a process used to determine the order and timing (chronology) of events using information in tree rings (dendro- refers to trees)
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Secondary growth pt.2
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 stormsâcalled cyclones in the Indian Ocean and typhoons in the Pacificâthat 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 Asiaâs rainfall, as well as the regionâs 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 âmonsoon,â 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 cultivationâe.g., in Java (Indonesia) and western Luzon (the Philippines)âin 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 worldâs 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 AsiaâCelebes (Sulawesi), the Moluccas, and the Lesser Sunda Islandsâconstitutes 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 regionâs 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 Java
Endochondral ossification (contâd) Secondary ossification center appears in the epiphysis (shortly after birth); no bone collar During remodeling, cavity is produced that fills with bone marrow 2 areas of cartilage remain Articular Epiphyseal (responsible for bone growth in length) Closure of plates is complete by age 20 Once closed, growth in length is stopped
What is a Plant Cell? Plant cells are eukaryotic cells that vary in several fundamental factors from other eukaryotic organisms. Both plant and animal cells contain a nucleus along with similar organelles. One of the distinctive aspects of a plant cell is the presence of a cell wall outside the cell membrane. Plant Cell Structure Just like different organs within the body, plant cell structure includes various components known as cell organelles that perform different functions to sustain itself. These organelles include: Cell Wall It is a rigid layer which is composed of polysaccharides cellulose, pectin and hemicellulose. It is located outside the cell membrane. It also comprises glycoproteins and polymers such as lignin, cutin, or suberin. The primary function of the cell wall is to protect and provide structural support to the cell. The plant cell wall is also involved in protecting the cell against mechanical stress and providing form and structure to the cell. It also filters the molecules passing in and out of it. The formation of the cell wall is guided by microtubules. It consists of three layers, namely, primary, secondary and the middle lamella. The primary cell wall is formed by cellulose laid down by enzymes. Cell membrane It is the semi-permeable membrane that is present within the cell wall. It is composed of a thin layer of protein and fat. The cell membrane plays an important role in regulating the entry and exit of specific substances within the cell. For instance, cell membrane keeps toxins from entering inside, while nutrients and essential minerals are transported across. Nucleus The nucleus is a membrane-bound structure that is present only in eukaryotic cells. The vital function of a nucleus is to store DNA or hereditary information required for cell division, metabolism and growth. 1. Nucleolus: It manufactures cellsâ protein-producing structures and ribosomes. 2. Nucleopore: Nuclear membrane is perforated with holes called nucleopore that allow proteins and nucleic acids to pass through. Plastids They are membrane-bound organelles that have their own DNA. They are necessary to store starch and to carry out the process of photosynthesis. It is also used in the synthesis of many molecules, which form the building blocks of the cell. Some of the vital types of plastids and their functions are stated below: Leucoplasts They are found in the non-photosynthetic tissue of plants. They are used for the storage of protein, lipid and starch. Chromoplasts They are heterogeneous, colored plastid which is responsible for pigment synthesis and for storage in photosynthetic eukaryotic organisms. Chromoplasts have red-, orange- and yellow-colored pigments which provide color to all ripe fruits and flowers. Central Vacuole It occupies around 30% of the cellâs volume in a mature plant cell. Tonoplast is a membrane that surrounds the central vacuole. The vital function of the central vacuole apart from storage is to sustain turgor pressure against the cell wall. The central vacuole consists of cell sap. It is a mixture of salts, enzymes and other substances. Golgi Apparatus They are found in all eukaryotic cells, which are involved in distributing synthesized macromolecules to various parts of the cell. Ribosomes They are the smallest membrane-bound organelles which comprise RNA and protein. They are the sites for protein synthesis, hence, also referred to as the protein factories of the cell. Mitochondria They are the double-membraned organelles found in the cytoplasm of all eukaryotic cells. They provide energy by breaking down carbohydrate and sugar molecules, hence they are also referred to as the âPowerhouse of the cell.â Lysosome Lysosomes are called suicidal bags as they hold digestive enzymes in an enclosed membrane. They perform the function of cellular waste disposal by digesting worn-out organelles, food particles and foreign bodies in the cell. In plants, the role of lysosomes is undertaken by the vacuoles. Chloroplasts It is an elongated organelle enclosed by phospholipid membrane. The chloroplast is shaped like a disc and the stroma is the fluid within the chloroplast that comprises a circular DNA. Each chloroplast contains a green colored pigment called chlorophyll required for the process of photosynthesis. The chlorophyll absorbs light energy from the sun and uses it to transform carbon dioxide and water into glucose. Structure of Chloroplast Chloroplasts are found in all higher plants. It is oval or biconvex, found within the mesophyll of the plant cell. The size of the chloroplast usually varies between 4-6 ”m in diameter and 1-3 ”m in thickness. They are double-membrane organelle with the presence of outer, inner and intermembrane space. There are two distinct regions present inside a chloroplast known as the grana and stroma. âą Grana are made up of stacks of disc-shaped structures known as thylakoids or lamellae. The granum of the chloroplast consists of chlorophyll pigments and are the functional units of chloroplasts. âą Stroma is the homogenous matrix which contains grana and is similar to the cytoplasm in cells in which all the organelles are embedded. Stroma also contains various enzymes, DNA, ribosomes, and other substances. Stroma lamellae function by connecting the stacks of thylakoid sacs or grana. The chloroplast structure consists of the following parts: Membrane Envelope It comprises inner and outer lipid bilayer membranes. The inner membrane separates the stroma from the intermembrane space. Intermembrane Space The space between inner and outer membranes. Thylakoid System (Lamellae) The system is suspended in the stroma. It is a collection of membranous sacs called thylakoids or lamellae. The green colored pigments called chlorophyll are found in the thylakoid membranes. It is the sight for the process of light-dependent reactions of the photosynthesis process. The thylakoids are arranged in stacks known as grana and each granum contains around 10-20 thylakoids. Stroma It is a colorless, alkaline, aqueous, protein-rich fluid present within the inner membrane of the chloroplast present surrounding the grana. Grana Stack of lamellae in plastids is known as grana. These are the sites of conversion of light energy into chemical energy. Chlorophyll It is a green photosynthetic pigment that helps in the process of photosynthesis. Functions of Chloroplast Following are the important chloroplast functions: âą The most important function of the chloroplast is to synthesize food by the process of photosynthesis. âą Absorbs light energy and converts it into chemical energy. âą Chloroplast has a structure called chlorophyll which functions by trapping the solar energy and is used for the synthesis of food in all green plants. âą Produces NADPH and molecular oxygen (O 2 ) by photolysis of water. âą Produces ATP â Adenosine triphosphate by the process of photosynthesis. âą The carbon dioxide (CO2) obtained from the air is used to generate carbon and sugar during the Calvin Cycle or dark reaction of photosynthesis. Mitochondria âMitochondria are membrane-bound organelles present in the cytoplasm of all eukaryotic cells, that produce adenosine triphosphate (ATP), the main energy molecule used by the cell.â What are Mitochondria? Popularly known as the âPowerhouse of the cell,â mitochondria (singular: mitochondrion) are a double membrane-bound organelle found in most eukaryotic organisms. They are found inside the cytoplasm and essentially function as the cellâs âdigestive system.â They play a major role in breaking down nutrients and generating energy-rich molecules for the cell. Many of the biochemical reactions involved in cellular respiration take place within the mitochondria. The term âmitochondrionâ is derived from the Greek words âmitosâ and âchondrionâ which means âthreadâ and âgranules-likeâ, respectively. It was first described by a German pathologist named Richard Altmann in the year 1890. Structure of Mitochondria âą The mitochondrion is a double-membraned, rod-shaped structure found in both plant and animal cell. âą Its size ranges from 0.5 to 1.0 micrometers in diameter. âą The structure comprises an outer membrane, an inner membrane, and a gel-like material called the matrix. âą The outer membrane and the inner membrane are made of proteins and phospholipid layers separated by the intermembrane space. âą The outer membrane covers the surface of the mitochondrion and has a large number of special proteins known as porins. Cristae The inner membrane of mitochondria is rather complex in structure. It has many folds that form a layered structure called cristae, and this helps in increasing the surface area inside the organelle. The cristae and the proteins of the inner membrane aid in the production of ATP molecules. The inner mitochondrial membrane is strictly permeable only to oxygen and ATP molecules. A number of chemical reactions take place within the inner membrane of mitochondria. Mitochondrial Matrix The mitochondrial matrix is a viscous fluid that contains a mixture of enzymes and proteins. It also comprises ribosomes, inorganic ions, mitochondrial DNA, nucleotide cofactors, and organic molecules. The enzymes present in the matrix play an important role in the synthesis of ATP molecules. Functions of Mitochondria The most important function of mitochondria is to produce energy through the process of oxidative phosphorylation. It is also involved in the following process: 1. Regulates the metabolic activity of the cell 2. Promotes the growth of new cells and cell multiplication 3. Helps in detoxifying ammonia in the liver cells 4. Plays an important role in apoptosis or programmed cell death 5. Responsible for building certain parts of the blood and various hormones like testosterone and estrogen 6. Helps in maintaining an adequate concentration of calcium ions within the compartments of the cell 7. It is also involved in various cellular activities like cellular differentiation, cell signaling, cell senescence, controlling the cell cycle and in cell growth. Disorders Associated with Mitochondria Any irregularity in the way mitochondria function can directly affect human health, but often, it is difficult to identify because symptoms differ from person to person. Disorders of the mitochondria can be quite severe; in some cases, they can even cause an organ to fail.
Nutrition Notes Nutrition- study of how your body uses food Process by which body uses nutrients How you look and feel Resist diseases and illness How you perform physically and mentally Nutrients: substances in food your body needs to grow, repair and supply energy to your body cells 6 Classes of Nutrients 1.Carbohydrates: 1 gram= 4 calories 2. Protein: 1 gram- 4 calories 3. Fats: 1 gram= 9 calories 4.Water 5. Vitamins 6. Minerals Calorie: measurement of energy in food Metabolism: Rate at which body burns energy(calories) Hunger: physical drive to eat Appetite: pshycological desire for food What influences your food choices: Foods you like Health Reasons Family and Culture Time & Money Advertising Emotions Friends Social Media: Modeling Nutrients Carbohydrates: your bodyâs main source of energy sugars/starches in food 45%-65% of diet #1 source of energy Simple: sugars converted to glucose= energy (fruits, dairy, honey, some manufactured foods) Complex: sugars linked together (starches) (grains, bread, pasta, beans, vegetables) Fiber: tough, indigestible carbohydrates Cleans our digestive system Prevents some types of cancer Prevents heart disease (fruits, vegetables, whole grains,nuts) 2. Protein: growth and repair of body tissues Made up of chemicals called âamino acidsâ Basic building material of all body cells (muscles, bones, skin, internal organs) Secondary source of energy protein(hemoglobin) attaches to oxygen in blood Functions as hormones regulating body functions 10-15% of diet *Body uses 20 Amino Acids found in food ( body produces 11 and 9 must come from diet) Essential amino acids: 9 amino acids body doesn't produce Complete Amino Acids: foods that contain all 9 essential amino acids ( animal products) Incomplete Amino Acids: food products that do not contain all 9 essential amino acids. Fats 15-25% of diet Secondary source of energy Blood clotting Controlling inflammation Maintains healthy skin/hair absorb /transport fat soluble vitamins Regulates body temperature Types of Fat Unsaturated: âgoodâ fat Liquid at room temperature Can help fight heart disease (veg oil, nuts) Saturated: âbadâ fat Solid at room temp Clogs arteries Causes strokes, heart attack, diabetes (animal products, meat, dairy) Cholesterol: waxy like fat substance found in meat products HDL: good type of cholesterol Body creates(liver) Creates cell wall, hormones, and vit D LDL: bad cholesterol- found in foods (clogs arteries) 4. Trans Fat: âone of the worst type of fatsâ Formed by a process called âhydrogenationâ: adding Hydrogen molecules to unsaturated fats to make it more solid and resistant to chemical change. Vitamins A vitamin is a chemical compound that is needed in small amounts for the human body to work correctly. Vitamins are classified as either fat soluble (vitamins A, D, E and K) or water soluble (vitamins B and C). This difference between the two groups is very important. It determines how each vitamin acts within the body. The fat soluble vitamins are soluble in lipids (fats). Fat soluble vitamins can be stored in our body Water soluble vitamins must be taken every day Human body produces some amounts of Vitamin D & K
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 plantâs lifestyle differs from an animalâs. 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 â List three structures that are present in plant cells but not in animal cells. â Compare the plasma membrane, the primary cell wall, and the secondary cell wall. â Explain the role of the central vacuole. â Describe the roles of plastids in the life of a plant. â 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 cellâs 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 cellâs volume and can push all of the other organelles into a thin layer against the plasma membrane. When water is plentiful, it fills a plantâs 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 âpale green,â and plastos, meaning âformedâ 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
Create a review game for 9th grade biology students using the following topics Levels of Organization in an ecosystem- population, community, ecosystem, biome, biosphere Abiotic and Biotic Factors Differences between Food chains and food webs Trophic Levels Producers vs Consumers, Autotrophs vs. Heterotrophs Effects of Greenhouse gases and their effects on global systems. Biome examples Photosynthesis vs cellular respiration Types of Consumers Ecological Pyramids 10% rule Cycles of Matter/ Nutrient Cycles- Water Cycle, Carbon Cycle, Nitrogen Cycle, Phosphorus Cycle (note on the diagrams⊠the bigger the arrow, the larger amount of matter that moves through the cycle from that point to the next. Macromolecules- Carbohydrates, Lipids, Proteins, Nucleic Acids Nitrogen fixation Denitrification Eutrophication The usable form on nitrogen for plants is nitrate Population density and distribution-random, dispersed and clumped Birth rate and death rate Survivorship curves- Type I, II, and III Density dependent factors Density independent factors Exponential growth- J curve = unlimited resources, no limiting factors Logistical Growth-S curve= limiting factors, carrying capacity Symbiotic Relationships- Competition, predation, Herbivory, mutualism, parasitism, commensalism What is an invasive species? Why might countries limit certain species to coming into a new country or area? What is mycorrhizal? Succession- Primary vs Secondary Pioneer Species Climax community Biodiversity Climate change
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 DesirĂ©e Nieuwenhuis | NatĂ lia 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 âonset of puberty and adiposity/obesityâ, âonset of pubertyâ, âchildren with obesityâ, âadipose tissueâ, âchildhood obesityâ, âadiposityâ, âobesityâ, âadipokine(s)â, âHPG axisâ, âadipokines ovary/ova- riesâ, or âadipokines testesâ, 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