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Chapter 8 – Photosynthesis Photosynthesis and Energy o Plants make their own food o They use the energy provided by the sun to take a simple gas, carbon dioxide, join it to a carbohydrate (a sugar) and then energize that sugar thus transforming it into food o From Plants, a Great Bounty for Animals A by-product of photosynthesis is oxygen Plants break water molecules apart In doing so, they use electrons and protons from H2O but they leave behind oxygen molecules (O2) Energy comes from the sun and then, in photosynthesis, is stored in plants in the complex molecules carbohydrates Thus stored, these carbohydrates can be broken down and used The breakdown of food ends with the cellular respiration that provides ATP o Up and Down the Energy Hill Again Photosynthesis and cellular respiration are trips up and down the energy hill Respiration – from more stored energy (in food) to less, as the food was broken down to produce ATP Photosynthesis is a trip up the energy hill Electrons are removed from water, boosted to a more energetic state by the power of sunlight and then brought together with a sugar and carbon dioxide resulting in an energy rich sugar Photosynthesis is the process by which certain groups of organisms capture energy from sunlight and convert this solar energy into chemical energy that is initially stored in a carbohydrate The Components of Photosynthesis o Starts with the absorption of sunlight by leaves o Absorption: light is taken in by the leaves, the leaves capture only a portion of the light that falls on them o Photosynthesis is driven by part of the visible light spectrum – mainly by blue and red light of certain wavelengths o This is why plants are green – they strongly scatter the green portion of the visible light spectrum o Where in the Plant Does Photosynthesis Occur? Leaf – one layer of outer (epidermal) cells at the top, another layer of epidermal cells at the bottom and several layers of mesophyll cells in between Chloroplasts are the sites of photosynthesis The stomata seen in the figure are the microscopic pores that let carbon dioxide pass into leaves and water vapor pass out of them Chloroplasts – organelles within plant and algae cells that are the sites of photosynthesis The chloroplast has outer and inner membranes at its periphery The interior of the chloroplast – there is a network of chloroplast membranes, active in photosynthesis called thylakoids (thylakoids often stack on top of one another creating structures called grana) Thylakoids are immersed in the liquid material of the chloroplast, the stroma Thylakoid membranes and the stroma are the places where all the steps of photosynthesis occur The thylakoid membranes are where photosynthesis start, with the absorption of sunlight Any compound that strongly absorbs certain visible wavelengths of sunlight is called a pigment Thylakoid membranes contain a pigment, called chlorophyll a that can be defined as the primary pigment active in plant photosynthesis Chlorophyll a is aided by several substances known as accessory pigments, which do just what their name implies: aid chlorophyll a in absorbing energetic rays from the sun after which they pass the absorbed energy along o There Are Two Essential Stages in Photosynthesis Two main stages of photosynthesis The first stage (the photo) – the power of sunlight will do two things – string water of electrons and then boost these electrons to a higher energy level The first stage ends when the original, energized electrons get attached to a mobile electron carrier called NADP+ The second stage (the synthesis) the electrons come together with carbon dioxide and a sugar This sugar produces a high energy sugar – meaning food This second stage takes place in the stroma or the chloroplast The first stage of photosynthesis obviously depends directly on the sunlight – these steps are sometimes referred to as light reactions The second set of steps is referred to as the Calvin Cycle o The Working Units of the Light Reactions Light reactions take place within a set of working units Photosystem – an organized complex of molecules with a thylakoid membrane that, in photosynthesis, collects solar energy and transforms it into chemical energy First, there is a group of a few hundred pigment molecules that serve to absorb the sunlight Majority of these molecules serve only as “antennae” that absorb energy from the sun and pass it on The center of the antennae system – the reaction center – a pair of special chlorophyll a molecules and associated compounds that first receive the solar energy from photosystem pigments and then transform this solar energy into chemical energy Includes a molecule that could be thought of as the first recipient of the absorbed solar energy the “primary electron acceptor” The absorbed energy is then passed on to the pair of chlorophull a molecules in a reaction center Electrons from this pair are “moved” in a couple of ways Physically these elements are transferred to the primary electron receptor Also movement up the energy hill The energy from sunlight is pumping the electrons up the hill in photosynthesis o Energy Transfer in Photosynthesis Works Through Redox Reactions Any time one substance loses (or “donates”) electrons to another it is said to have been oxidized The substance that gained electrons is said to have been reduced This is a redox reaction Many of the photosynthetic steps are redox reactions The movement of electrons with the reaction center – from the chlorphyll a molecules to the primary electron acceptor – is a redox reaction The chlorophyll molecules are oxidized by the primary electron acceptor Electrons are passed on by means of one electron carrier oxidizing another The electrons are moving both up and down the energy hill With the energy supplied by the sun, electrons are pumped up a couple of formidable energy gradients only to come partway back down them as they “seek” their lowest energy state They are releasing energy as they fall Stage 1: The Steps of the Light Reactions o First step is that solar energy, collected by photosystem II’s antennae molecules arrives at the reaction center o This energy then gives a boost to an electron in the reaction center in the two ways noted: the electron physicall moves to another part of the reaction center complex, the primary electron acceptor, and it is also pumped up the energy hill o The reaction center chlorophyll has lost an electron o That loss leaves an energy hole in this chlorophyll making it an oxidizing agent o A special enzyme in the reaction center splits water molecules that lie within the thylakoid compartment o These water molecules are now being oxidized which means they are losing electrons o These electrons travel to the reaction center, where they will be the next electrons in line for an energy boost o A Chain of Redox Reactions and Another Boost from the Sun After arriving at the primary electron acceptor they fall back down the energy hill as they are transferred through a series of electron transport molecules each oxidizing its predecessor They arrive at photosystem I which includes a slightly different kind fo reaction center This center is also receiving solar energy and uses it to boost electrons to a higher energy state Electrons are transferred down the second energy hill – until they are received by the electron carrier NADP+ In accepting electrons NADP+ becomes reduced to NADPH an electron carrier that ferries the electrons into the next stage of photosynthesis This second stage is the Calvin cycle which will yield the high energy sugar that is the essential product of photosynthesis o The Physical Movement of Electrons in the Light Reactions Electrons have physically moved through the chloroplast Started out in the water of the thylakoid, through the thylakoid membrane and ended up in the stroma attached to NADPH What Makes the Light Reactions so Important? o Two momentous things that have taken place within these steps: the splitting of water and the transformation of solar energy to chemical energy o The Splitting of Water: Electrons and Oxygen The splitting of water provides the traveling electrons Oxygen accounts for 21 percent of the Earth’s atmosphere In the water splitting hydrogen atoms are removed from H2O while oxygen is left behind When two liberated oxygen atoms come together in the thylakoid compartment the result is O2 – the form of oxygen that exists in the atmosphere o The Transformation of Solar Energy to Chemical Energy Transformation of solar energy to chemical energy is the second major feat that takes place Energy of the sunlight moves to a chlorophyll molecule in a reaction center, it boosts an electron there from what is known as a ground state to an excited state This electron is moving farther out from the nucleus of an atom One fate is for the electrons to drop back down to the original state in the process releasing as heat then energy they have absorbed Another possible fate is for falling electrons to release part of their energy as light in the process known as fluorescence In photosynthesis the energized electrons are transferred to a different molecule and they don’t fall back to their ground state They are passed on a redox reaction Electrons are taken from water, boosted to a higher energy state by the sun and transferred to other molecules o Production of ATP A primary function of the fall of electrons between photosystems II and I is a release of energy that is used for the production of ATP ATP will be used to power the reactions that are coming up in the second stage of photosynthesis The suns power has provided energy that is stored in two forms – ATP and the energetic electrons in NADPH Stage 2: The Calvin Cycle o The energy captured in the light reactions will be used o It will power a process by which carbon dioxide taken from the atmosphere – will be joined to a sugar, with the resulting product energized, thus creating a carbohydrate o Energized Sugar Comes from a Cycle of Reactions Calvin Cycle – or the C3 cycle – is the set of steps in photosynthesis in which energetic electrons are brought together with carbon dioxide and a sugar to produce an energetic carbohydrate Synthesis of photosynthesis – a bring together of elements The first stems can be thought of as a process of fixation – of a gas being incorporated into an organic molecule Carbon dioxide, which comes in through the leaves of the plant, is being fixed into the starting sugar, which is called RuBP It is the way life builds itself up with materials that lie outside of life The next reactions in the cycle are the energizing steps of the process Low energy sugar receives the energetic products of the light reactions The RuBP derivatives first interact with ATP and then receive energetic electrons from NADPH A relatively low energy sugar has been energized from which position it can now serve as food The energized sugar – G3P – is the essential product of photosynthesis One molecule of it is netted with each turn of the Calvin cycle The remainder of the cycle can be thought of as a preparation of molecules for another trip around the cycle o The Ultimate Product of Photosynthesis G3P can be turned into many things Put two G3P molecules together and you get the more familiar six carbon sugar glucose The ultimate product of photosynthesis is the whole plant Photorespiration and the C4 Pathway o A glitch in this process is called photorespiration and it takes place within the Calvin cycle o Cycle begins with carbon dioxide being joined to a low energy sugar o What brings these two compounds together is an enzyme called rubisco o It is extremely powerful but slow moving and prone to making errors o Rubisco is the bridge between the living world and the nonliving world o It is the molecule that allows plants to capture the carbon atoms that are the building blocks of all living things o One of the problems is its speed – rubsico can only manage 3 reactions per second as opposed to the average of 1000 o Because rubisco works so slowly, every plant must have a lot of it as a result rubisco is the most abundant protein on Earth o Rubsico has a problem in even binding with the right substance - it can bind to oxygen as well as to carbon dioxide o Rubisco may fix up to one molecule of O2 for every three of the carbon dioxide o Photorespiration is especially likely to take place when temperatures rise because heat prompts the stomata on leaves to close to preserve water o The effect of closed stomata is not only that water is kept in but that CO2 is kept out o With a deficit of CO2 rubisco tends to bind more frequently with oxygen o The result is plants that don’t grow as much o Photorespiration: a process in which the enzyme rubisco undercuts carbon fixation in photosynthesis by binding with oxygen instead of with carbon dioxide o Rubisco: can be defined as an enzyme that allows organisms to incorporate atmospheric carbon dioxide into their own sugars during the process of photosynthesis o C4 Photosynthesis is a form of photosynthesis in which carbon dioxide is first fixed to a four carbon molecule and then transferred to special cells in which the Calvin cycle is undertaken, bundle sheath cells o The C4 Pathway Is Not Always Advantageous C4 Pathway is known to be employed most notably in some grasses, and in corn, sugar can and sorghum – vastly more C3 than C4 plants C4 fixation does not confer an across the board advantage It has a cost, which is the expenditure of ATP in shuttling carbon dioxide to the bundle sheath cells C4 fixation is advantageous only where the weather is warm enough to bring about a significant increase in photorespiration Another Photosynthetic Variation: CAM Plants o When plants live in climates that are not just warm but dry, a large part of their survival comes down to retaining water o Photosynthesis, however, works against water retention o When CO2 can pass in, water vapor can pass out o Plans we call succulents – like cacti – have a solution for this problem – close their stomata during the day and open them at night o The plants then carry out C4 metabolism at night but only up to a point o They fix carbon dioxide into an initial four carbon molecule and then stand pat o The CO2 stays banked in them awaiting the energy of the next days sun o CAM Photosynthesis is a form of photosynthesis, undertaken by plants in hot, dry climates, in which carbon fixation takes place at night and the Calvin cycle occurs during the day o CAM is an acronym: crassulacean acid metabolism