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Plant physiology · Plant nutrition

Photosynthesis

Picture a soybean field in Iowa in July, or the oak trees along a Texas Hill Country road: almost all the organic matter and almost all the oxygen on the planet have the same origin, a reaction that takes place in a green leaf when light falls on it. Understanding how that reaction works, step by step, means understanding the basis of almost all life on Earth. By the end of this page you will be able to explain why a plant in a sunny greenhouse can still stop growing faster.

The overall equation and the pigments

Place a sprig of Elodea in illuminated water and you will see a continuous string of bubbles rising, which stops when you cover the beaker: which gas is being produced, and which inorganic materials does all the mass the plant gains as it grows come from?

$$6\ \text{CO}_2 + 6\ \text{H}_2\text{O} + \text{light energy} \rightarrow \text{C}_6\text{H}_{12}\text{O}_6 + 6\ \text{O}_2$$ A chloroplast with its stacks of thylakoids receives light, carbon dioxide and water, and releases oxygen as bubbles and a glucose molecule. light chloroplast 6 CO₂ 6 H₂O 6 O₂ glucose C₆H₁₂O₆

Why photosynthesis matters

Photosynthesis matters for three reasons. Matter: it turns inorganic carbon ($\text{CO}_2$) into organic molecules that build the bodies of producers and, through food chains, of every consumer. Energy: it converts light energy into chemical energy stored in glucose, which feeds nearly all ecosystems. Atmosphere: it releases $\text{O}_2$ for aerobic respiration and removes $\text{CO}_2$ from the air.

Gas exchange: stomata

$\text{CO}_2$ enters the leaf through tiny pores in the epidermis called stomata (singular: stoma), each opened and closed by two guard cells. Water vapor and $\text{O}_2$ leave through the same pores. The sugar made in the leaf travels out through the phloem in the veins to roots, fruits and growing tissues.

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Write the overall chemical equation for photosynthesis and identify the reactants, products, and energy source of the process.

Write the overall chemical equation for photosynthesis and identify the reactants, products, and energy source of the process.

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Which pigment captures light during photosynthesis and in which cell organelle is it located? Explain why leaves are green and what happens to this color in the fall in…

Which pigment captures light during photosynthesis and in which cell organelle is it located? Explain why leaves are green and what happens to this color in the fall in deciduous species.

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State and explain three reasons why photosynthesis is essential for life on Earth, considering matter, energy, and the atmosphere.

State and explain three reasons why photosynthesis is essential for life on Earth, considering matter, energy, and the atmosphere.

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Name the main photosynthetic pigments (chlorophylls a and b, carotenoids) and explain what the absorption spectrum is: which colors of light they use and which they…

Name the main photosynthetic pigments (chlorophylls a and b, carotenoids) and explain what the absorption spectrum is: which colors of light they use and which they reflect, relating this to the color we see.

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The chloroplast: thylakoids and stroma

A leaf cell can hold dozens of chloroplasts, yet glucose is not made just anywhere inside them. Where exactly do the two halves of photosynthesis happen?

Section of a chloroplast showing its double membrane, the thylakoids stacked in grana and joined by lamellae, where the light-dependent stage takes place, and the stroma with the Calvin cycle enzymes. Chloroplast outer membrane inner membrane thylakoid enzymes (fix CO₂) grana (stacked thylakoids) light-dependent reactions stroma Calvin cycle
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Explain where photosynthesis occurs inside the chloroplast: what the thylakoids and the grana are and what the stroma is, indicating which phase of the process occurs in…

Explain where photosynthesis occurs inside the chloroplast: what the thylakoids and the grana are and what the stroma is, indicating which phase of the process occurs in each compartment.

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Distinguish between the light-dependent stage and the light-independent stage (Calvin cycle) of photosynthesis: in which part of the chloroplast each occurs, what…

Distinguish between the light-dependent stage and the light-independent stage (Calvin cycle) of photosynthesis: in which part of the chloroplast each occurs, what substances enter, and what is produced in each stage.

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The light-dependent reactions: splitting of water

For decades it was believed that the oxygen given off by plants came from $\text{CO}_2$, until labeling water molecules with heavy isotopes showed that all the gas came from the water absorbed: how can solar energy manage to break such a stable chemical bond?

$$2\ \text{H}_2\text{O} \xrightarrow{\text{light}} 4\ \text{H}^+ + 4\ e^- + \text{O}_2$$ Photolysis of water in the thylakoid membrane: the light captured by chlorophyll splits two water molecules inside the thylakoid and produces four protons, which stay in the lumen, four electrons, which leave towards ATP and NADPH synthesis, and one oxygen molecule, which is released. stroma inside the thylakoid (lumen) thylakoid membrane chlorophyll light 2 H₂O photolysis 4 e⁻ towards ATP and NADPH H⁺ H⁺ H⁺ H⁺ 4 H⁺ O₂ → is released 2 H₂O light 4 H⁺ + 4 e⁻ + O₂
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Which molecule does the oxygen released by photosynthesis come from: $\text{CO}2$ or water? Explain how this was tested using labeled isotopes ($\text{O-18}$) and why…

Which molecule does the oxygen released by photosynthesis come from: $\text{CO}_2$ or water? Explain how this was tested using labeled isotopes ($\text{O-18}$) and why the result was decisive.

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Explain what the photolysis of water is and which three products it generates (electrons, protons and oxygen), indicating what the plant uses the first two for.

Explain what the photolysis of water is and which three products it generates (electrons, protons and oxygen), indicating what the plant uses the first two for.

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In 1882, T. W. Engelmann illuminated a filament of the green alga Spirogyra* with a visible light spectrum obtained using a prism, using motile aerotactic bacteria to…

In 1882, T. W. Engelmann illuminated a filament of the green alga Spirogyra with a visible light spectrum obtained using a prism, using motile aerotactic bacteria to map oxygen production.

  1. a) Identify the independent variable, the dependent variable, and the biosensor in this experimental design, justifying the biological basis of the bacterial movement.
  2. b) Predict along which wavelengths of the spectrum ($400\text{--}700\ \text{nm}$) the densest bacterial aggregates were observed and justify why this distribution correlates mainly with the absorption spectrum of chlorophylls and cannot be explained by that of accessory carotenoids alone.
  3. c) Imagine that the experiment is repeated replacing Spirogyra with a deep benthic cyanobacterium equipped with phycobilins (pigments with optimal absorption between $500$ and $600\ \text{nm}$). Predict how the pattern of bacterial aggregation would change and deduce the ecological advantage of these pigments in aquatic habitats.
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The Calvin cycle: fixing carbon

The sugar in a Thanksgiving pumpkin pie started as invisible gas in the air. How does a plant turn $\text{CO}_2$ into a solid molecule?

Circular diagram of the Calvin cycle in a clockwise direction: CO₂ enters and is fixed by RuBisCO, ATP and NADPH are supplied, G3P leaves and ribulose bisphosphate (RuBP) is regenerated. CO₂ enters (fixed by RuBisCO) ATP, NADPH G3P (leaves) RuBP is regenerated Calvin cycle (stroma)

Back to the start: the bubbles from Elodea are $\text{O}_2$ from the splitting of water, and the plant's added mass comes from $\text{CO}_2$ and water.

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The overall equations for photosynthesis and cellular respiration appear symmetrical. Explain why, despite this, they are not reverse processes: compare organelles,…

The overall equations for photosynthesis and cellular respiration appear symmetrical. Explain why, despite this, they are not reverse processes: compare organelles, mechanisms, and the net energy direction of each.

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The diagram shows the exchange of matter in a leaf during photosynthesis. a) Indicate which substance is carried by the arrow marked "?" that exits through the vein and…

The diagram shows the exchange of matter in a leaf during photosynthesis. a) Indicate which substance is carried by the arrow marked "?" that exits through the vein and to which parts of the plant it travels. b) Write the overall equation for the process and relate each arrow to the reactants or products. c) Name the epidermal structure through which $\text{CO}_2$ enters.

light H₂O ? CO₂ O₂
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Limiting factors: light, CO₂, temperature and C4/CAM strategies

Light compensation point

A plant respires all the time but photosynthesizes only in light. The light compensation point is the light intensity at which photosynthesis equals respiration, so net $\text{O}_2$ exchange is zero. Below it the plant loses more carbon than it gains and must live off stored starch; understory plants have low compensation points because they are adapted to dim light.

Photorespiration

RuBisCO can also bind $\text{O}_2$ instead of $\text{CO}_2$. This wasteful reaction, photorespiration, releases carbon and wastes ATP, and it gets worse in hot, dry air when stomata close and $\text{O}_2$ builds up in the leaf. Lowering the $\text{O}_2$ around a C3 plant raises its net photosynthesis.

Starch test and the Keeling curve

Iodine turns blue-black in the presence of starch, so a decolorized (boiled in ethanol to remove chlorophyll, which would hide the color) leaf stained with iodine shows where photosynthesis made sugar: green areas of a variegated leaf turn blue-black, white areas do not.

Atmospheric $\text{CO}_2$ at Mauna Loa rises each year but also oscillates seasonally: it falls in the Northern Hemisphere growing season, when its large land masses of plants take up $\text{CO}_2$, and rises in fall and winter when respiration and decay exceed photosynthesis. The Southern Hemisphere has much less land vegetation, so it does not cancel this signal.

Productivity of the ocean

Phytoplankton are less than 1% of photosynthetic biomass but about half of net primary production because their cells divide in days and are eaten quickly (high $P/B$), while trees store carbon for decades. The ocean has a second puzzle: why are warm tropical seas, with plenty of light and heat, often so unproductive? The reason is not P/B but nutrients. In these seas a permanent thermocline (a sharp temperature boundary) keeps warm surface water from mixing with deeper, nutrient-rich water, so nutrients, not light or temperature, are the limiting factor for phytoplankton growth.

When a plant's output is measured in the laboratory under ever more powerful lamps, the photosynthetic rate at first rises in direct proportion but then levels off on a horizontal plateau: why does supplying more light energy stop speeding up the reaction?

Graph of photosynthetic rate against light intensity: it rises almost in a straight line at low light, gradually curves and levels off on a plateau where another factor (CO₂ or temperature) limits photosynthesis. light is limiting plateau: another factor is limiting (CO₂, temperature…) light intensity rate of photosynthesis
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Name three environmental factors that influence the photosynthetic rate (light intensity, $\text{CO}2$ concentration, and temperature) and explain briefly how each one…

Name three environmental factors that influence the photosynthetic rate (light intensity, $\text{CO}_2$ concentration, and temperature) and explain briefly how each one acts when it is scarce or excessive.

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What is the light compensation point? Explain what happens when photosynthesis equals respiration, what happens to a plant that lives for weeks below that point (where…

What is the light compensation point? Explain what happens when photosynthesis equals respiration, what happens to a plant that lives for weeks below that point (where does it get energy from?) and why understory plants have lower compensation points than full-sun ones.

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Greenhouses enrich the air with $\text{CO}2$ to increase the harvest. Explain why it works by using the law of the minimum (the limiting factor), whether the increase in…

Greenhouses enrich the air with $\text{CO}_2$ to increase the harvest. Explain why it works by using the law of the minimum (the limiting factor), whether the increase in production that can be achieved is unlimited, and which other factors become limiting once $\text{CO}_2$ is corrected.

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A variegated leaf has white zones without chlorophyll. After a few hours of light, it is decolorized and stained with iodine. Predict which zones will stain blue-black,…

A variegated leaf has white zones without chlorophyll. After a few hours of light, it is decolorized and stained with iodine. Predict which zones will stain blue-black, explain what the result demonstrates (what does iodine detect?) and why the chlorophyll must be removed from the leaf beforehand.

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The concentration of atmospheric $\text{CO}2$ rises every year, but with a seasonal oscillation (the "saw teeth" of the Keeling curve). Explain what causes this annual…

The concentration of atmospheric $\text{CO}_2$ rises every year, but with a seasonal oscillation (the "saw teeth" of the Keeling curve). Explain what causes this annual oscillation, why it mainly reflects the photosynthesis of the Northern Hemisphere, and what it says about the scale of the biosphere's activity.

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A commercial tomato greenhouse in central California installs supplemental LED lighting to increase winter production. When irradiance is increased from $400$ to…

A commercial tomato greenhouse in central California installs supplemental LED lighting to increase winter production. When irradiance is increased from $400$ to $800\ \mu\text{mol}\times\text{m}^{-2}\times\text{s}^{-1}$, the $\text{CO}_2$ fixation rate only increases by $4\,\%$, reaching an unexpected plateau.

  1. a) Justify at the physiological and molecular level why, beyond a certain light intensity, the photosynthetic rate saturates even though more photons are available.
  2. b) Formulate a hypothesis regarding what alternative environmental factor is limiting the process in the greenhouse and design an experiment with controlled variables to test it.
  3. c) Reasonably predict how the light saturation curve and the maximum photosynthetic rate will change if the greenhouse atmosphere is enriched with gaseous $\text{CO}_2$.
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In an agronomic trial in a semiarid area subjected to a heat wave ($38\ ^\circ\text{C}$) and summer drought, three adjacent plots are monitored: wheat (C3), sorghum…

In an agronomic trial in a semiarid area subjected to a heat wave ($38\ ^\circ\text{C}$) and summer drought, three adjacent plots are monitored: wheat (C3), sorghum (C4), and prickly pear (Opuntia, CAM).

  1. a) Vacuolar acidity (malate concentration) and stomatal opening are measured at 03:00 in the morning and at 14:00 at midday. Predict which crop will show maximum acidity at 03:00 and justify how this adaptation allows it to survive with minimal water expenditure.
  2. b) During the central hours of the day, wheat drastically reduces its net carbon assimilation rate to near-zero values, while sorghum maintains a very high positive net rate. Justify this contrast at the enzymatic level (RuBisCO versus PEP carboxylase) in relation to stomatal closure and photorespiration.
  3. c) In an experimental scenario with artificially enriched $\text{CO}_2$ but without water deficit, predict which of the two crops (C3 or C4) will experience a greater percentage increase in its photosynthetic rate and reason why.
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The Keeling curve shows atmospheric $\text{CO}2$ concentration at Mauna Loa since 1958, combining a continuous upward trend with an annual periodic "sawtooth"…

The Keeling curve shows atmospheric $\text{CO}_2$ concentration at Mauna Loa since 1958, combining a continuous upward trend with an annual periodic "sawtooth" oscillation (maximum in May, minimum in October).

  1. a) Interpret the global biological cause of the annual oscillation and reason why the $\text{CO}_2$ minimum is recorded at the beginning of the Northern Hemisphere autumn (October) and not at the summer solstice, and why the Southern Hemisphere seasons do not neutralize this oscillation.
  2. b) At the South Pole measuring station, the underlying upward trend is identical, but the annual oscillation has an almost negligible amplitude (less than $1\ \text{ppm}$ compared to $\sim 6\ \text{ppm}$ in Hawaii). Justify this discrepancy biogeographically.
  3. c) Knowing that terrestrial and oceanic biological sinks currently absorb about $50\,\%$ of anthropogenic $\text{CO}_2$, predict and justify how the slope of the Keeling curve would change if heat stress were to transform boreal forests from net carbon sinks into net carbon emitters.
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The enzyme RuBisCO is bifunctional: it catalyzes both the carboxylation (photosynthesis) and the oxygenation of ribulose-1.5-bisphosphate (photorespiration). In a plant…

The enzyme RuBisCO is bifunctional: it catalyzes both the carboxylation (photosynthesis) and the oxygenation of ribulose-1.5-bisphosphate (photorespiration). In a plant physiology experiment, identical soybean plants (C3 plant) are grown at $30\ ^\circ\text{C}$ in two chambers with identical light and the same concentration of $\text{CO}_2$ ($0.04\,\%$), but varying oxygen: Chamber 1 ($21\,\%$ $\text{O}_2$) and Chamber 2 ($2\,\%$ $\text{O}_2$).

  1. a) Predict in which chamber the net photosynthetic rate and biomass gain will be higher. Justify your deduction by explaining the energy and carbon balance of photorespiration.
  2. b) If the temperature in both chambers is raised from $30\ ^\circ\text{C}$ to $40\ ^\circ\text{C}$, the photosynthetic yield in Chamber 1 drops drastically, whereas in Chamber 2 it remains almost constant. Justify this result considering the enzymatic affinity of RuBisCO and the relative solubility of both gases as water warms up.
  3. c) Biotechnology aims to introduce C4 pathways into C3 crops such as wheat or rice. Argue whether this modification would provide a clear adaptive advantage in cold and temperate latitudes or if, on the contrary, it could reduce their yield compared to unmodified varieties.
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Although marine phytoplankton represents less than $1\,\%$ of the global photosynthetic biomass, it is responsible for nearly $50\,\%$ of the net primary production of…

Although marine phytoplankton represents less than $1\,\%$ of the global photosynthetic biomass, it is responsible for nearly $50\,\%$ of the net primary production of the biosphere.

  1. a) Resolve this apparent ecological paradox by comparing the residence time of organic matter and the turnover rate (production/biomass ratio, $P/B$) of marine microalgae versus the trees of a temperate forest.
  2. b) In tropical seas where solar radiation and temperature are optimal year-round, photosynthetic productivity is typically extraordinarily low. Justify the role of the permanent thermocline in this phenomenon and predict the impact of ocean surface warming on the extent of these oligotrophic areas.
  3. c) Explain how the ocean's "biological carbon pump" works and predict the consequences of a phytoplankton decline on both global warming and the higher trophic levels of the pelagic marine food web.
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Investigating a limiting factor with data

The question "does photosynthesis increase as light increases?" requires defining what is changed, what is measured and what is kept constant. A high school lab design can analyze simulated data on the net oxygen production of an aquatic plant at controlled temperature and CO₂.

Relative light Replicate 1 Replicate 2 Replicate 3
0 −2 −1 −3
1 3 4 5
2 7 8 9
3 8 9 10

The values are arbitrary units of net O₂ exchange per minute, not real measurements. Plot the means in a spreadsheet, with the independent variable (light) on the x-axis and the dependent variable (net $\text{O}_2$ exchange) on the y-axis.

Here is a quick design for the Elodea experiment: hypothesis (the closer the lamp, the more bubbles per minute), independent variable (distance to the lamp), dependent variable (bubbles per minute), controls (same temperature using a water bath, same $\text{CO}_2$ and plant sample, plus a no-plant beaker). One limitation: counting bubbles is imprecise because bubble size varies.

Experimental set-up: a lamp illuminates an aquatic plant submerged in a beaker of water, with a thermometer and an oxygen sensor connected to a display; the light is changed, the O₂ exchange is measured, and species, biomass, CO₂, temperature and duration are kept constant. Light Temperature species and biomass CO₂ O₂ O₂ exchange duration changed measured kept constant
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Design an experiment with an aquatic plant (Elodea) to measure how photosynthesis varies with the distance to a lamp: hypothesis, independent variable, how the rate is…

Design an experiment with an aquatic plant (Elodea) to measure how photosynthesis varies with the distance to a lamp: hypothesis, independent variable, how the rate is measured (bubbles of $\text{O}_2$ per minute), two necessary controls, and one limitation of the method.

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Summary

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