title: Chloroplast

text: All chloroplasts have at least three membrane systems—the outer chloroplast membrane, the inner chloroplast membrane, and the thylakoid system. Chloroplasts that are the product of secondary endosymbiosis may have additional membranes surrounding these three. Inside the outer and inner chloroplast membranes is the chloroplast stroma, a semi-gel-like fluid that makes up much of a chloroplast's volume, and in which the thylakoid system floats.


title: Chloroplast

text: There are some common misconceptions about the outer and inner chloroplast membranes. The fact that chloroplasts are surrounded by a double membrane is often cited as evidence that they are the descendants of endosymbiotic cyanobacteria. This is often interpreted as meaning the outer chloroplast membrane is the product of the host's cell membrane infolding to form a vesicle to surround the ancestral cyanobacterium—which is not true—both chloroplast membranes are homologous to the cyanobacterium's original double membranes.


title: Chloroplast

text: The chloroplast double membrane is also often compared to the mitochondrial double membrane. This is not a valid comparison—the inner mitochondria membrane is used to run proton pumps and carry out oxidative phosphorylation across to generate ATP energy. The only chloroplast structure that can considered analogous to it is the internal thylakoid system. Even so, in terms of "in-out", the direction of chloroplast H ion flow is in the opposite direction compared to oxidative phosphorylation in mitochondria. In addition, in terms of function, the inner chloroplast membrane, which regulates metabolite passage and synthesizes some materials, has no counterpart in the mitochondrion.


title: Chloroplast

text: The outer chloroplast membrane is a semi-porous membrane that small molecules and ions can easily diffuse across. However, it is not permeable to larger proteins, so chloroplast polypeptides being synthesized in the cell cytoplasm must be transported across the outer chloroplast membrane by the TOC complex, or "translocon on the outer chloroplast" membrane.


title: Chloroplast

text: The chloroplast membranes sometimes protrude out into the cytoplasm, forming a stromule, or stroma-containing tubule. Stromules are very rare in chloroplasts, and are much more common in other plastids like chromoplasts and amyloplasts in petals and roots, respectively. They may exist to increase the chloroplast's surface area for cross-membrane transport, because they are often branched and tangled with the endoplasmic reticulum. When they were first observed in 1962, some plant biologists dismissed the structures as artifactual, claiming that stromules were just oddly shaped chloroplasts with constricted regions or dividing chloroplasts. However, there is a growing body of evidence that stromules are functional, integral features of plant cell plastids, not merely artifacts.


title: Chloroplast

text: Usually, a thin intermembrane space about 10–20 nanometers thick exists between the outer and inner chloroplast membranes.


title: Chloroplast

text: Glaucophyte algal chloroplasts have a peptidoglycan layer between the chloroplast membranes. It corresponds to the peptidoglycan cell wall of their cyanobacterial ancestors, which is located between their two cell membranes. These chloroplasts are called "muroplasts" (from Latin ""mura"", meaning "wall"). Other chloroplasts were assumed to have lost the cyanobacterial wall, leaving an intermembrane space between the two chloroplast envelope membranes, but has since been found also in moss, lycophytes and ferns.


title: Chloroplast

text: The inner chloroplast membrane borders the stroma and regulates passage of materials in and out of the chloroplast. After passing through the TOC complex in the outer chloroplast membrane, polypeptides must pass through the TIC complex "(translocon on the inner chloroplast membrane)" which is located in the inner chloroplast membrane.


title: Chloroplast

text: In addition to regulating the passage of materials, the inner chloroplast membrane is where fatty acids, lipids, and carotenoids are synthesized.


title: Chloroplast

text: Some chloroplasts contain a structure called the chloroplast peripheral reticulum. It is often found in the chloroplasts of plants, though it has also been found in some angiosperms, and even some gymnosperms. The chloroplast peripheral reticulum consists of a maze of membranous tubes and vesicles continuous with the inner chloroplast membrane that extends into the internal stromal fluid of the chloroplast. Its purpose is thought to be to increase the chloroplast's surface area for cross-membrane transport between its stroma and the cell cytoplasm. The small vesicles sometimes observed may serve as transport vesicles to shuttle stuff between the thylakoids and intermembrane space.


title: Chloroplast

text: The protein-rich, alkaline, aqueous fluid within the inner chloroplast membrane and outside of the thylakoid space is called the stroma, which corresponds to the cytosol of the original cyanobacterium. Nucleoids of chloroplast DNA, chloroplast ribosomes, the thylakoid system with plastoglobuli, starch granules, and many proteins can be found floating around in it. The Calvin cycle, which fixes CO into G3P takes place in the stroma.


title: Chloroplast

text: Chloroplasts have their own ribosomes, which they use to synthesize a small fraction of their proteins. Chloroplast ribosomes are about two-thirds the size of cytoplasmic ribosomes (around 17 nm vs 25 nm). They take mRNAs transcribed from the chloroplast DNA and translate them into protein. While similar to bacterial ribosomes, chloroplast translation is more complex than in bacteria, so chloroplast ribosomes include some chloroplast-unique features.


title: Chloroplast

text: Small subunit ribosomal RNAs in several Chlorophyta and euglenid chloroplasts lack motifs for Shine-Dalgarno sequence recognition, which is considered essential for translation initiation in most chloroplasts and prokaryotes. Such loss is also rarely observed in other plastids and prokaryotes. An additional 4.5S rRNA with homology to the 3' tail of 23S is found in "higher" plants.


title: Chloroplast

text: Plastoglobuli (singular "plastoglobulus", sometimes spelled "plastoglobule(s)"), are spherical bubbles of lipids and proteins about 45–60 nanometers across. They are surrounded by a lipid monolayer. Plastoglobuli are found in all chloroplasts, but become more common when the chloroplast is under oxidative stress, or when it ages and transitions into a gerontoplast. Plastoglobuli also exhibit a greater size variation under these conditions. They are also common in etioplasts, but decrease in number as the etioplasts mature into chloroplasts.


title: Chloroplast

text: Plastoglubuli contain both structural proteins and enzymes involved in lipid synthesis and metabolism. They contain many types of lipids including plastoquinone, vitamin E, carotenoids and chlorophylls.


title: Chloroplast

text: Plastoglobuli were once thought to be free-floating in the stroma, but it is now thought that they are permanently attached either to a thylakoid or to another plastoglobulus attached to a thylakoid, a configuration that allows a plastoglobulus to exchange its contents with the thylakoid network. In normal green chloroplasts, the vast majority of plastoglobuli occur singularly, attached directly to their parent thylakoid. In old or stressed chloroplasts, plastoglobuli tend to occur in linked groups or chains, still always anchored to a thylakoid.


title: Chloroplast

text: Plastoglobuli form when a bubble appears between the layers of the lipid bilayer of the thylakoid membrane, or bud from existing plastoglubuli—though they never detach and float off into the stroma. Practically all plastoglobuli form on or near the highly curved edges of the thylakoid disks or sheets. They are also more common on stromal thylakoids than on granal ones.


title: Chloroplast

text: Starch granules are very common in chloroplasts, typically taking up 15% of the organelle's volume, though in some other plastids like amyloplasts, they can be big enough to distort the shape of the organelle. Starch granules are simply accumulations of starch in the stroma, and are not bounded by a membrane.


title: Chloroplast

text: Starch granules appear and grow throughout the day, as the chloroplast synthesizes sugars, and are consumed at night to fuel respiration and continue sugar export into the phloem, though in mature chloroplasts, it is rare for a starch granule to be completely consumed or for a new granule to accumulate.


title: Chloroplast

text: Starch granules vary in composition and location across different chloroplast lineages. In red algae, starch granules are found in the cytoplasm rather than in the chloroplast. In plants, mesophyll chloroplasts, which do not synthesize sugars, lack starch granules.


title: Chloroplast

text: The chloroplast stroma contains many proteins, though the most common and important is RuBisCO, which is probably also the most abundant protein on the planet. RuBisCO is the enzyme that fixes CO into sugar molecules. In plants, RuBisCO is abundant in all chloroplasts, though in plants, it is confined to the bundle sheath chloroplasts, where the Calvin cycle is carried out in plants.


title: Chloroplast

text: The chloroplasts of some hornworts and algae contain structures called pyrenoids. They are not found in higher plants. Pyrenoids are roughly spherical and highly refractive bodies which are a site of starch accumulation in plants that contain them. They consist of a matrix opaque to electrons, surrounded by two hemispherical starch plates. The starch is accumulated as the pyrenoids mature. In algae with carbon concentrating mechanisms, the enzyme RuBisCO is found in the pyrenoids. Starch can also accumulate around the pyrenoids when CO is scarce. Pyrenoids can divide to form new pyrenoids, or be produced "de novo".


title: Chloroplast

text: Thylakoids (sometimes spelled "thylakoïds"), are small interconnected sacks which contain the membranes that the light reactions of photosynthesis take place on. The word "thylakoid" comes from the Greek word "thylakos" which means "sack".


title: Chloroplast

text: Suspended within the chloroplast stroma is the thylakoid system, a highly dynamic collection of membranous sacks called thylakoids where chlorophyll is found and the light reactions of photosynthesis happen.


title: Chloroplast

text: In most vascular plant chloroplasts, the thylakoids are arranged in stacks called grana, though in certain plant chloroplasts and some algal chloroplasts, the thylakoids are free floating.


title: Chloroplast

text: Using a light microscope, it is just barely possible to see tiny green granules—which were named grana. With electron microscopy, it became possible to see the thylakoid system in more detail, revealing it to consist of stacks of flat thylakoids which made up the grana, and long interconnecting stromal thylakoids which linked different grana.


title: Chloroplast

text: In the transmission electron microscope, thylakoid membranes appear as alternating light-and-dark bands, 8.5 nanometers thick.


title: Chloroplast

text: For a long time, the three-dimensional structure of the thylakoid membrane system had been unknown or disputed. Many models have been proposed, the most prevalent being the helical model, in which granum stacks of thylakoids are wrapped by helical stromal thylakoids. Another model known as the 'bifurcation model', which was based on the first electron tomography study of plant thylakoid membranes, depicts the stromal membranes as wide lamellar sheets perpendicular to the grana columns which bifurcates into multiple parallel discs forming the granum-stroma assembly. The helical model was supported by several additional works, but ultimately it was determined in 2019 that features from both the helical and bifurcation models are consolidated by newly discovered left-handed helical membrane junctions. Likely for ease, the thylakoid system is still commonly depicted by older "hub and spoke" models where the grana are connected to each other by tubes of stromal thylakoids.


title: Chloroplast

text: Grana consist of a stacks of flattened circular granal thylakoids that resemble pancakes. Each granum can contain anywhere from two to a hundred thylakoids, though grana with 10–20 thylakoids are most common. Wrapped around the grana are multiple parallel right-handed helical stromal thylakoids, also known as frets or lamellar thylakoids. The helices ascend at an angle of ~20°, connecting to each granal thylakoid at a bridge-like slit junction.


title: Chloroplast

text: The stroma lamellae extend as large sheets perpendicular to the grana columns. These sheets are connected to the right-handed helices either directly or through bifurcations that form left-handed helical membrane surfaces. The left-handed helical surfaces have a similar tilt angle to the right-handed helices (~20°), but ¼ the pitch. Approximately 4 left-handed helical junctions are present per granum, resulting in a pitch-balanced array of right- and left-handed helical membrane surfaces of different radii and pitch that consolidate the network with minimal surface and bending energies. While different parts of the thylakoid system contain different membrane proteins, the thylakoid membranes are continuous and the thylakoid space they enclose form a single continuous labyrinth.


title: Chloroplast

text: Embedded in the thylakoid membranes are important protein complexes which carry out the light reactions of photosynthesis. Photosystem II and photosystem I contain light-harvesting complexes with chlorophyll and carotenoids that absorb light energy and use it to energize electrons. Molecules in the thylakoid membrane use the energized electrons to pump hydrogen ions into the thylakoid space, decreasing the pH and turning it acidic. ATP synthase is a large protein complex that harnesses the concentration gradient of the hydrogen ions in the thylakoid space to generate ATP energy as the hydrogen ions flow back out into the stroma—much like a dam turbine.


title: Chloroplast

text: There are two types of thylakoids—granal thylakoids, which are arranged in grana, and stromal thylakoids, which are in contact with the stroma. Granal thylakoids are pancake-shaped circular disks about 300–600 nanometers in diameter. Stromal thylakoids are helicoid sheets that spiral around grana. The flat tops and bottoms of granal thylakoids contain only the relatively flat photosystem II protein complex. This allows them to stack tightly, forming grana with many layers of tightly appressed membrane, called granal membrane, increasing stability and surface area for light capture.


title: Chloroplast

text: In contrast, photosystem I and ATP synthase are large protein complexes which jut out into the stroma. They can't fit in the appressed granal membranes, and so are found in the stromal thylakoid membrane—the edges of the granal thylakoid disks and the stromal thylakoids. These large protein complexes may act as spacers between the sheets of stromal thylakoids.


title: Chloroplast

text: The number of thylakoids and the total thylakoid area of a chloroplast is influenced by light exposure. Shaded chloroplasts contain larger and more grana with more thylakoid membrane area than chloroplasts exposed to bright light, which have smaller and fewer grana and less thylakoid area. Thylakoid extent can change within minutes of light exposure or removal.


title: Chloroplast

text: Inside the photosystems embedded in chloroplast thylakoid membranes are various photosynthetic pigments, which absorb and transfer light energy. The types of pigments found are different in various groups of chloroplasts, and are responsible for a wide variety of chloroplast colorations.


title: Chloroplast

text: Chlorophyll "a" is found in all chloroplasts, as well as their cyanobacterial ancestors. Chlorophyll "a" is a blue-green pigment partially responsible for giving most cyanobacteria and chloroplasts their color. Other forms of chlorophyll exist, such as the accessory pigments chlorophyll "b", chlorophyll "c", chlorophyll "d", and chlorophyll "f".


title: Chloroplast

text: Chlorophyll "b" is an olive green pigment found only in the chloroplasts of plants, green algae, any secondary chloroplasts obtained through the secondary endosymbiosis of a green alga, and a few cyanobacteria. It is the chlorophylls "a" and "b" together that make most plant and green algal chloroplasts green.


title: Chloroplast

text: Chlorophyll "c" is mainly found in secondary endosymbiotic chloroplasts that originated from a red alga, although it is not found in chloroplasts of red algae themselves. Chlorophyll "c" is also found in some green algae and cyanobacteria.


title: Chloroplast

text: In addition to chlorophylls, another group of yellow–orange pigments called carotenoids are also found in the photosystems. There are about thirty photosynthetic carotenoids. They help transfer and dissipate excess energy, and their bright colors sometimes override the chlorophyll green, like during the fall, when the leaves of some land plants change color. β-carotene is a bright red-orange carotenoid found in nearly all chloroplasts, like chlorophyll "a". Xanthophylls, especially the orange-red zeaxanthin, are also common. Many other forms of carotenoids exist that are only found in certain groups of chloroplasts.


title: Chloroplast

text: Phycobilins are a third group of pigments found in cyanobacteria, and glaucophyte, red algal, and cryptophyte chloroplasts. Phycobilins come in all colors, though phycoerytherin is one of the pigments that makes many red algae red. Phycobilins often organize into relatively large protein complexes about 40 nanometers across called phycobilisomes. Like photosystem I and ATP synthase, phycobilisomes jut into the stroma, preventing thylakoid stacking in red algal chloroplasts. Cryptophyte chloroplasts and some cyanobacteria don't have their phycobilin pigments organized into phycobilisomes, and keep them in their thylakoid space instead.


title: Chloroplast

text: To fix carbon dioxide into sugar molecules in the process of photosynthesis, chloroplasts use an enzyme called RuBisCO. RuBisCO has trouble distinguishing between carbon dioxide and oxygen, so at high oxygen concentrations, RuBisCO starts accidentally adding oxygen to sugar precursors. This has the result of ATP energy being wasted and being released, all with no sugar being produced. This is a big problem, since O is produced by the initial light reactions of photosynthesis, causing issues down the line in the Calvin cycle which uses RuBisCO.


title: Chloroplast

text: plants evolved a way to solve this—by spatially separating the light reactions and the Calvin cycle. The light reactions, which store light energy in ATP and NADPH, are done in the mesophyll cells of a leaf. The Calvin cycle, which uses the stored energy to make sugar using RuBisCO, is done in the bundle sheath cells, a layer of cells surrounding a vein in a leaf.


title: Chloroplast

text: As a result, chloroplasts in mesophyll cells and bundle sheath cells are specialized for each stage of photosynthesis. In mesophyll cells, chloroplasts are specialized for the light reactions, so they lack RuBisCO, and have normal grana and thylakoids, which they use to make ATP and NADPH, as well as oxygen. They store in a four-carbon compound, which is why the process is called " photosynthesis". The four-carbon compound is then transported to the bundle sheath chloroplasts, where it drops off and returns to the mesophyll. Bundle sheath chloroplasts do not carry out the light reactions, preventing oxygen from building up in them and disrupting RuBisCO activity. Because of this, they lack thylakoids organized into grana stacks—though bundle sheath chloroplasts still have free-floating thylakoids in the stroma where they still carry out cyclic electron flow, a light-driven method of synthesizing ATP to power the Calvin cycle without generating oxygen. They lack photosystem II, and only have photosystem I—the only protein complex needed for cyclic electron flow. Because the job of bundle sheath chloroplasts is to carry out the Calvin cycle and make sugar, they often contain large starch grains.


title: Chloroplast

text: Both types of chloroplast contain large amounts of chloroplast peripheral reticulum, which they use to get more surface area to transport stuff in and out of them. Mesophyll chloroplasts have a little more peripheral reticulum than bundle sheath chloroplasts.


title: Chloroplast

text: Not all cells in a multicellular plant contain chloroplasts. All green parts of a plant contain chloroplasts—the chloroplasts, or more specifically, the chlorophyll in them are what make the photosynthetic parts of a plant green. The plant cells which contain chloroplasts are usually parenchyma cells, though chloroplasts can also be found in collenchyma tissue. A plant cell which contains chloroplasts is known as a chlorenchyma cell. A typical chlorenchyma cell of a land plant contains about 10 to 100 chloroplasts.


title: Chloroplast

text: In some plants such as cacti, chloroplasts are found in the stems, though in most plants, chloroplasts are concentrated in the leaves. One square millimeter of leaf tissue can contain half a million chloroplasts. Within a leaf, chloroplasts are mainly found in the mesophyll layers of a leaf, and the guard cells of stomata. Palisade mesophyll cells can contain 30–70 chloroplasts per cell, while stomatal guard cells contain only around 8–15 per cell, as well as much less chlorophyll. Chloroplasts can also be found in the bundle sheath cells of a leaf, especially in C plants, which carry out the Calvin cycle in their bundle sheath cells. They are often absent from the epidermis of a leaf.


title: Chloroplast

text: The chloroplasts of plant and algal cells can orient themselves to best suit the available light. In low-light conditions, they will spread out in a sheet—maximizing the surface area to absorb light. Under intense light, they will seek shelter by aligning in vertical columns along the plant cell's cell wall or turning sideways so that light strikes them edge-on. This reduces exposure and protects them from photooxidative damage. This ability to distribute chloroplasts so that they can take shelter behind each other or spread out may be the reason why land plants evolved to have many small chloroplasts instead of a few big ones.


title: Chloroplast

text: Chloroplast movement is considered one of the most closely regulated stimulus-response systems that can be found in plants. Mitochondria have also been observed to follow chloroplasts as they move.


title: Chloroplast

text: In higher plants, chloroplast movement is run by phototropins, blue light photoreceptors also responsible for plant phototropism. In some algae, mosses, ferns, and flowering plants, chloroplast movement is influenced by red light in addition to blue light, though very long red wavelengths inhibit movement rather than speeding it up. Blue light generally causes chloroplasts to seek shelter, while red light draws them out to maximize light absorption.


title: Chloroplast

text: Studies of "Vallisneria gigantea", an aquatic flowering plant, have shown that chloroplasts can get moving within five minutes of light exposure, though they don't initially show any net directionality. They may move along microfilament tracks, and the fact that the microfilament mesh changes shape to form a honeycomb structure surrounding the chloroplasts after they have moved suggests that microfilaments may help to anchor chloroplasts in place.


title: Chloroplast

text: Unlike most epidermal cells, the guard cells of plant stomata contain relatively well-developed chloroplasts. However, exactly what they do is controversial.


title: Chloroplast

text: Plants lack specialized immune cells—all plant cells participate in the plant immune response. Chloroplasts, along with the nucleus, cell membrane, and endoplasmic reticulum, are key players in pathogen defense. Due to its role in a plant cell's immune response, pathogens frequently target the chloroplast.


title: Chloroplast

text: Plants have two main immune responses—the hypersensitive response, in which infected cells seal themselves off and undergo programmed cell death, and systemic acquired resistance, where infected cells release signals warning the rest of the plant of a pathogen's presence.


title: Chloroplast

text: Chloroplasts stimulate both responses by purposely damaging their photosynthetic system, producing reactive oxygen species. High levels of reactive oxygen species will cause the hypersensitive response. The reactive oxygen species also directly kill any pathogens within the cell. Lower levels of reactive oxygen species initiate systemic acquired resistance, triggering defense-molecule production in the rest of the plant.


title: Chloroplast

text: In some plants, chloroplasts are known to move closer to the infection site and the nucleus during an infection.


title: Chloroplast

text: Chloroplasts can serve as cellular sensors. After detecting stress in a cell, which might be due to a pathogen, chloroplasts begin producing molecules like salicylic acid, jasmonic acid, nitric oxide and reactive oxygen species which can serve as defense-signals. As cellular signals, reactive oxygen species are unstable molecules, so they probably don't leave the chloroplast, but instead pass on their signal to an unknown second messenger molecule. All these molecules initiate retrograde signaling—signals from the chloroplast that regulate gene expression in the nucleus.


title: Chloroplast

text: In addition to defense signaling, chloroplasts, with the help of the peroxisomes, help synthesize an important defense molecule, jasmonate. Chloroplasts synthesize all the fatty acids in a plant cell—linoleic acid, a fatty acid, is a precursor to jasmonate.


title: Chloroplast

text: One of the main functions of the chloroplast is its role in photosynthesis, the process by which light is transformed into chemical energy, to subsequently produce food in the form of sugars. Water (HO) and carbon dioxide (CO) are used in photosynthesis, and sugar and oxygen (O) is made, using light energy. Photosynthesis is divided into two stages—the light reactions, where water is split to produce oxygen, and the dark reactions, or Calvin cycle, which builds sugar molecules from carbon dioxide. The two phases are linked by the energy carriers adenosine triphosphate (ATP) and nicotinamide adenine dinucleotide phosphate (NADP).


title: Chloroplast

text: The light reactions take place on the thylakoid membranes. They take light energy and store it in NADPH, a form of NADP, and ATP to fuel the dark reactions.


title: Chloroplast

text: ATP is the phosphorylated version of adenosine diphosphate (ADP), which stores energy in a cell and powers most cellular activities. ATP is the energized form, while ADP is the (partially) depleted form. NADP is an electron carrier which ferries high energy electrons. In the light reactions, it gets reduced, meaning it picks up electrons, becoming NADPH.


title: Chloroplast

text: Like mitochondria, chloroplasts use the potential energy stored in an H, or hydrogen ion gradient to generate ATP energy. The two photosystems capture light energy to energize electrons taken from water, and release them down an electron transport chain. The molecules between the photosystems harness the electrons' energy to pump hydrogen ions into the thylakoid space, creating a concentration gradient, with more hydrogen ions (up to a thousand times as many) inside the thylakoid system than in the stroma. The hydrogen ions in the thylakoid space then diffuse back down their concentration gradient, flowing back out into the stroma through ATP synthase. ATP synthase uses the energy from the flowing hydrogen ions to phosphorylate adenosine diphosphate into adenosine triphosphate, or ATP. Because chloroplast ATP synthase projects out into the stroma, the ATP is synthesized there, in position to be used in the dark reactions.


title: Chloroplast

text: Electrons are often removed from the electron transport chains to charge NADP with electrons, reducing it to NADPH. Like ATP synthase, ferredoxin-NADP reductase, the enzyme that reduces NADP, releases the NADPH it makes into the stroma, right where it is needed for the dark reactions.


title: Chloroplast

text: Because NADP reduction removes electrons from the electron transport chains, they must be replaced—the job of photosystem II, which splits water molecules (HO) to obtain the electrons from its hydrogen atoms.


title: Chloroplast

text: While photosystem II photolyzes water to obtain and energize new electrons, photosystem I simply reenergizes depleted electrons at the end of an electron transport chain. Normally, the reenergized electrons are taken by NADP, though sometimes they can flow back down more H-pumping electron transport chains to transport more hydrogen ions into the thylakoid space to generate more ATP. This is termed cyclic photophosphorylation because the electrons are recycled. Cyclic photophosphorylation is common in plants, which need more ATP than NADPH.


title: Chloroplast

text: The Calvin cycle, also known as the dark reactions, is a series of biochemical reactions that fixes CO into G3P sugar molecules and uses the energy and electrons from the ATP and NADPH made in the light reactions. The Calvin cycle takes place in the stroma of the chloroplast.


title: Chloroplast

text: While named ""the dark reactions"", in most plants, they take place in the light, since the dark reactions are dependent on the products of the light reactions.


title: Chloroplast

text: The Calvin cycle starts by using the enzyme RuBisCO to fix CO into five-carbon Ribulose bisphosphate (RuBP) molecules. The result is unstable six-carbon molecules that immediately break down into three-carbon molecules called 3-phosphoglyceric acid, or 3-PGA.


title: Chloroplast

text: The ATP and NADPH made in the light reactions is used to convert the 3-PGA into glyceraldehyde-3-phosphate, or G3P sugar molecules. Most of the G3P molecules are recycled back into RuBP using energy from more ATP, but one out of every six produced leaves the cycle—the end product of the dark reactions.


title: Chloroplast

text: Glyceraldehyde-3-phosphate can double up to form larger sugar molecules like glucose and fructose. These molecules are processed, and from them, the still larger sucrose, a disaccharide commonly known as table sugar, is made, though this process takes place outside of the chloroplast, in the cytoplasm.


title: Chloroplast

text: Alternatively, glucose monomers in the chloroplast can be linked together to make starch, which accumulates into the starch grains found in the chloroplast.


title: Chloroplast

text: Under conditions such as high atmospheric CO concentrations, these starch grains may grow very large, distorting the grana and thylakoids. The starch granules displace the thylakoids, but leave them intact.


title: Chloroplast

text: Waterlogged roots can also cause starch buildup in the chloroplasts, possibly due to less sucrose being exported out of the chloroplast (or more accurately, the plant cell). This depletes a plant's free phosphate supply, which indirectly stimulates chloroplast starch synthesis.


title: Chloroplast

text: While linked to low photosynthesis rates, the starch grains themselves may not necessarily interfere significantly with the efficiency of photosynthesis, and might simply be a side effect of another photosynthesis-depressing factor.


title: Chloroplast

text: Photorespiration can occur when the oxygen concentration is too high. RuBisCO cannot distinguish between oxygen and carbon dioxide very well, so it can accidentally add O instead of CO to RuBP. This process reduces the efficiency of photosynthesis—it consumes ATP and oxygen, releases CO, and produces no sugar. It can waste up to half the carbon fixed by the Calvin cycle. Several mechanisms have evolved in different lineages that raise the carbon dioxide concentration relative to oxygen within the chloroplast, increasing the efficiency of photosynthesis. These mechanisms are called carbon dioxide concentrating mechanisms, or CCMs. These include Crassulacean acid metabolism, carbon fixation, and pyrenoids. Chloroplasts in plants are notable as they exhibit a distinct chloroplast dimorphism.


title: Chloroplast

text: Because of the H gradient across the thylakoid membrane, the interior of the thylakoid is acidic, with a pH around 4, while the stroma is slightly basic, with a pH of around 8.


title: Chloroplast

text: The optimal stroma pH for the Calvin cycle is 8.1, with the reaction nearly stopping when the pH falls below 7.3.


title: Chloroplast

text: CO in water can form carbonic acid, which can disturb the pH of isolated chloroplasts, interfering with photosynthesis, even though CO is used in photosynthesis. However, chloroplasts in living plant cells are not affected by this as much.


title: Chloroplast

text: Chloroplasts can pump K and H ions in and out of themselves using a poorly understood light-driven transport system.


title: Chloroplast

text: In the presence of light, the pH of the thylakoid lumen can drop up to 1.5 pH units, while the pH of the stroma can rise by nearly one pH unit.


title: Chloroplast

text: Chloroplasts alone make almost all of a plant cell's amino acids in their stroma except the sulfur-containing ones like cysteine and methionine. Cysteine is made in the chloroplast (the proplastid too) but it is also synthesized in the cytosol and mitochondria, probably because it has trouble crossing membranes to get to where it is needed. The chloroplast is known to make the precursors to methionine but it is unclear whether the organelle carries out the last leg of the pathway or if it happens in the cytosol.


title: Chloroplast

text: Chloroplasts make all of a cell's purines and pyrimidines—the nitrogenous bases found in DNA and RNA. They also convert nitrite (NO) into ammonia (NH) which supplies the plant with nitrogen to make its amino acids and nucleotides.


title: Chloroplast

text: The plastid is the site of diverse and complex lipid synthesis in plants. The carbon used to form the majority of the lipid is from acetyl-CoA, which is the decarboxylation product of pyruvate. Pyruvate may enter the plastid from the cytosol by passive diffusion through the membrane after production in glycolysis. Pyruvate is also made in the plastid from phosphoenolpyruvate, a metabolite made in the cytosol from pyruvate or PGA. Acetate in the cytosol is unavailable for lipid biosynthesis in the plastid. The typical length of fatty acids produced in the plastid are 16 or 18 carbons, with 0-3 cis double bonds.


title: Chloroplast

text: The biosynthesis of fatty acids from acetyl-CoA primarily requires two enzymes. Acetyl-CoA carboxylase creates malonyl-CoA, used in both the first step and the extension steps of synthesis. Fatty acid synthase (FAS) is a large complex of enzymes and cofactors including acyl carrier protein (ACP) which holds the acyl chain as it is synthesized. The initiation of synthesis begins with the condensation of malonyl-ACP with acetyl-CoA to produce ketobutyryl-ACP. 2 reductions involving the use of NADPH and one dehydration creates butyryl-ACP. Extension of the fatty acid comes from repeated cycles of malonyl-ACP condensation, reduction, and dehydration.


title: Chloroplast

text: Other lipids are derived from the methyl-erythritol phosphate (MEP) pathway and consist of gibberelins, sterols, abscisic acid, phytol, and innumerable secondary metabolites.


title: Chloroplast

text: Chloroplasts are a special type of a plant cell organelle called a plastid, though the two terms are sometimes used interchangeably. There are many other types of plastids, which carry out various functions. All chloroplasts in a plant are descended from undifferentiated proplastids found in the zygote, or fertilized egg. Proplastids are commonly found in an adult plant's apical meristems. Chloroplasts do not normally develop from proplastids in root tip meristems—instead, the formation of starch-storing amyloplasts is more common.


title: Chloroplast

text: In shoots, proplastids from shoot apical meristems can gradually develop into chloroplasts in photosynthetic leaf tissues as the leaf matures, if exposed to the required light. This process involves invaginations of the inner plastid membrane, forming sheets of membrane that project into the internal stroma. These membrane sheets then fold to form thylakoids and grana.


title: Chloroplast

text: If angiosperm shoots are not exposed to the required light for chloroplast formation, proplastids may develop into an etioplast stage before becoming chloroplasts. An etioplast is a plastid that lacks chlorophyll, and has inner membrane invaginations that form a lattice of tubes in their stroma, called a prolamellar body. While etioplasts lack chlorophyll, they have a yellow chlorophyll precursor stocked. Within a few minutes of light exposure, the prolamellar body begins to reorganize into stacks of thylakoids, and chlorophyll starts to be produced. This process, where the etioplast becomes a chloroplast, takes several hours. Gymnosperms do not require light to form chloroplasts.


title: Chloroplast

text: Light, however, does not guarantee that a proplastid will develop into a chloroplast. Whether a proplastid develops into a chloroplast some other kind of plastid is mostly controlled by the nucleus and is largely influenced by the kind of cell it resides in.


title: Chloroplast

text: Plastid differentiation is not permanent, in fact many interconversions are possible. Chloroplasts may be converted to chromoplasts, which are pigment-filled plastids responsible for the bright colors seen in flowers and ripe fruit. Starch storing amyloplasts can also be converted to chromoplasts, and it is possible for proplastids to develop straight into chromoplasts. Chromoplasts and amyloplasts can also become chloroplasts, like what happens when a carrot or a potato is illuminated. If a plant is injured, or something else causes a plant cell to revert to a meristematic state, chloroplasts and other plastids can turn back into proplastids. Chloroplast, amyloplast, chromoplast, proplast, etc., are not absolute states—intermediate forms are common.


title: Chloroplast

text: Most chloroplasts in a photosynthetic cell do not develop directly from proplastids or etioplasts. In fact, a typical shoot meristematic plant cell contains only 7–20 proplastids. These proplastids differentiate into chloroplasts, which divide to create the 30–70 chloroplasts found in a mature photosynthetic plant cell. If the cell divides, chloroplast division provides the additional chloroplasts to partition between the two daughter cells.


title: Chloroplast

text: In single-celled algae, chloroplast division is the only way new chloroplasts are formed. There is no proplastid differentiation—when an algal cell divides, its chloroplast divides along with it, and each daughter cell receives a mature chloroplast.


title: Chloroplast

text: Almost all chloroplasts in a cell divide, rather than a small group of rapidly dividing chloroplasts. Chloroplasts have no definite S-phase—their DNA replication is not synchronized or limited to that of their host cells.


title: Chloroplast

text: Much of what we know about chloroplast division comes from studying organisms like "Arabidopsis" and the red alga "Cyanidioschyzon merolæ".


title: Chloroplast

text: The division process starts when the proteins FtsZ1 and FtsZ2 assemble into filaments, and with the help of a protein ARC6, form a structure called a Z-ring within the chloroplast's stroma. The Min system manages the placement of the Z-ring, ensuring that the chloroplast is cleaved more or less evenly. The protein MinD prevents FtsZ from linking up and forming filaments. Another protein ARC3 may also be involved, but it is not very well understood. These proteins are active at the poles of the chloroplast, preventing Z-ring formation there, but near the center of the chloroplast, MinE inhibits them, allowing the Z-ring to form.


title: Chloroplast

text: Next, the two plastid-dividing rings, or PD rings form. The inner plastid-dividing ring is located in the inner side of the chloroplast's inner membrane, and is formed first. The outer plastid-dividing ring is found wrapped around the outer chloroplast membrane. It consists of filaments about 5 nanometers across, arranged in rows 6.4 nanometers apart, and shrinks to squeeze the chloroplast. This is when chloroplast constriction begins. In a few species like "Cyanidioschyzon merolæ", chloroplasts have a third plastid-dividing ring located in the chloroplast's intermembrane space.


title: Chloroplast

text: Late into the constriction phase, dynamin proteins assemble around the outer plastid-dividing ring, helping provide force to squeeze the chloroplast. Meanwhile, the Z-ring and the inner plastid-dividing ring break down. During this stage, the many chloroplast DNA plasmids floating around in the stroma are partitioned and distributed to the two forming daughter chloroplasts.


title: Chloroplast

text: Later, the dynamins migrate under the outer plastid dividing ring, into direct contact with the chloroplast's outer membrane, to cleave the chloroplast in two daughter chloroplasts.


title: Chloroplast

text: A remnant of the outer plastid dividing ring remains floating between the two daughter chloroplasts, and a remnant of the dynamin ring remains attached to one of the daughter chloroplasts.


title: Chloroplast

text: Of the five or six rings involved in chloroplast division, only the outer plastid-dividing ring is present for the entire constriction and division phase—while the Z-ring forms first, constriction does not begin until the outer plastid-dividing ring forms.


title: Chloroplast

text: In species of algae that contain a single chloroplast, regulation of chloroplast division is extremely important to ensure that each daughter cell receives a chloroplast—chloroplasts can't be made from scratch. In organisms like plants, whose cells contain multiple chloroplasts, coordination is looser and less important. It is likely that chloroplast and cell division are somewhat synchronized, though the mechanisms for it are mostly unknown.


title: Chloroplast

text: Light has been shown to be a requirement for chloroplast division. Chloroplasts can grow and progress through some of the constriction stages under poor quality green light, but are slow to complete division—they require exposure to bright white light to complete division. Spinach leaves grown under green light have been observed to contain many large dumbbell-shaped chloroplasts. Exposure to white light can stimulate these chloroplasts to divide and reduce the population of dumbbell-shaped chloroplasts.


title: Chloroplast

text: Like mitochondria, chloroplasts are usually inherited from a single parent. Biparental chloroplast inheritance—where plastid genes are inherited from both parent plants—occurs in very low levels in some flowering plants.


title: Chloroplast

text: Many mechanisms prevent biparental chloroplast DNA inheritance, including selective destruction of chloroplasts or their genes within the gamete or zygote, and chloroplasts from one parent being excluded from the embryo. Parental chloroplasts can be sorted so that only one type is present in each offspring.


title: Chloroplast

text: Gymnosperms, such as pine trees, mostly pass on chloroplasts paternally, while flowering plants often inherit chloroplasts maternally. Flowering plants were once thought to only inherit chloroplasts maternally. However, there are now many documented cases of angiosperms inheriting chloroplasts paternally.


title: Chloroplast

text: Angiosperms, which pass on chloroplasts maternally, have many ways to prevent paternal inheritance. Most of them produce sperm cells that do not contain any plastids. There are many other documented mechanisms that prevent paternal inheritance in these flowering plants, such as different rates of chloroplast replication within the embryo.


title: Chloroplast

text: Among angiosperms, paternal chloroplast inheritance is observed more often in hybrids than in offspring from parents of the same species. This suggests that incompatible hybrid genes might interfere with the mechanisms that prevent paternal inheritance.


title: Chloroplast

text: Recently, chloroplasts have caught attention by developers of genetically modified crops. Since, in most flowering plants, chloroplasts are not inherited from the male parent, transgenes in these plastids cannot be disseminated by pollen. This makes plastid transformation a valuable tool for the creation and cultivation of genetically modified plants that are biologically contained, thus posing significantly lower environmental risks. This biological containment strategy is therefore suitable for establishing the coexistence of conventional and organic agriculture. While the reliability of this mechanism has not yet been studied for all relevant crop species, recent results in tobacco plants are promising, showing a failed containment rate of transplastomic plants at 3 in 1,000,000.


title: My Secret Romance

text: My Secret Romance () is a 2017 South Korean television series starring Sung Hoon and Song Ji-eun. It aired on cable network OCN at 21:00 (KST) every Monday and Tuesday, from April 17 to May 30, 2017 for 14 episodes. It was also released on SK Telecom's "oksusu" mobile app.


title: My Secret Romance

text: "My Secret Romance" was selected in the 2017 Top Creator Audition held by KOCCA. The series is the first OCN drama to air on Monday and Tuesday, as well as the first romance-themed OCN drama.


title: My Secret Romance

text: Jin-wook and Yoo-mi meet at a Gangwon-do resort and get caught up in a series of misunderstandings and accidents. Yoo-mi is there to attend her mother's second wedding while Jin-wook is there working as a bellhop (a position given to him by his Chairman father to teach him responsibility). Yoo-mi is charmed by Jin-wook's sly and playful personality, and they unexpectedly spend the night together. However, Yoo-mi disappears in the morning, leaving Jin-wook feeling perplexed and insulted.


title: My Secret Romance

text: Three years later, the two meet again when Yoo-mi becomes a nutritionist at the company cafeteria where Jin-wook works. It seems that once a playboy Jin-wook, has given up his carefree life and works at the company owned by his father. He has feelings for Yoo-mi and has kept her bra-pad as a memory of the night that they spent together. At first, he is strict toward her, though it is only because he is looking for reasons to be near her. He later confesses this to her and agrees to wait for her.


title: My Secret Romance

text: During the three years, Yoo-mi's mother has had a second child, Dong-goo, who is now three years old. Upon looking at Dong-goo, Jin-wook's father, the Chairman, mistakes him as Yoo-mi's son, not half-brother. The Chairman tells Jin-wook, admonishing him for having a child out of wedlock with a nutritionist. After finding this out, Jin-wook believes that he has wronged Yoo-mi and warms up to her, seeming to accept Dong-goo as his son. The misunderstanding is eventually cleared up but Yoo-mi gets upset at Jin-wook, accusing him of being with her only because he thought he had a child with her. Then enters a pretty girl, Joo Hye-ri (Jung Da-sol) who is deemed a "suitable" wife for Jin-wook. Yoo-mi's writer friend, Jung Hyun-tae (Kim Jae-young), is shown to have a secret crush on her. With all drama revolving around main characters' mothers, Cha Jin-wook and Lee Yoo-mi finally manage to keep their love strong, while also resolving their personal problems with their parents.


title: My Secret Romance

text: The drama ends with Hye-ri "traveling together" with Hyun-tae and Yoo-mi and Jin-wook attending Yoo-mi's mother's re-wedding and Jin-wook saying "Today is D-day. Do you know how much I've waited for this day?" and Yoo-mi replying "Me too."


title: First Anglo-Afghan War

text: The First Anglo-Afghan War () was fought between the British Empire and the Emirate of Kabul from 1838 to 1842. The British initially successfully invaded the country taking sides in a succession dispute between emir Dost Mohammad (Barakzai) and former emir Shah Shujah (Durrani), whom they reinstalled upon occupying Kabul in August 1839. The main British Indian force occupied Kabul and endured harsh winters. The force and its camp followers were almost completely massacred during the 1842 retreat from Kabul.


title: First Anglo-Afghan War

text: The British then sent an "Army of Retribution" to Kabul to avenge the destruction of the previous forces. After recovering prisoners, they left Afghanistan by the end of the year. Dost Mohammed returned from exile in India to resume his rule.


title: First Anglo-Afghan War

text: It was one of the first major conflicts during the Great Game, the 19th century competition for power and influence in Central Asia between Britain and Russia.


title: First Anglo-Afghan War

text: The 19th century was a period of diplomatic competition between the British and Russian empires for spheres of influence in South Asia known as the "Great Game" to the British and the "Tournament of Shadows" to the Russians. With the exception of Emperor Paul who ordered an invasion of India in 1800 (which was cancelled after his assassination in 1801), no Russian tsar ever seriously considered invading India, but for most of the 19th century, Russia was viewed as "the enemy" in Britain; and any Russian advance into Central Asia, into what is now Kazakhstan, Turkmenistan, Kyrgyzstan, Uzbekistan and Tajikistan, was always assumed (in London) to be directed towards the conquest of India, as the American historian David Fromkin observed, "no matter how far-fetched" such an interpretation might be. In 1832, the First Reform Bill lowering the franchise requirements to vote and hold office in the United Kingdom was passed, which the ultra-conservative Emperor Nicholas I of Russia openly disapproved of, setting the stage for an Anglo-Russian "cold war", with many believing that Russian autocracy and British democracy were bound to clash. In 1837, Lord Palmerston and John Hobhouse, fearing the instability of Afghanistan, the Sindh, and the increasing power of the Sikh kingdom to the northwest, raised the spectre of a possible Russian invasion of British India through Afghanistan. The British tended to misunderstand the foreign policy of the Emperor Nicholas I as anti-British and intent upon an expansionary policy in Asia; whereas in fact though Nicholas disliked Britain as a liberal democratic state that he considered to be rather "strange", he always believed it was possible to reach an understanding with Britain on spheres of influence in Asia, believing that the essentially conservative nature of British society would retard the advent of liberalism. The main goal of Nicholas's foreign policy was not the conquest of Asia, but rather upholding the "status quo" in Europe, especially by co-operating with Prussia and Austria, and in isolating France, as Louis Philippe I, the King of the French was a man whom Nicholas hated as an "usurper". The duc d'Orleans had once been Nicholas's friend, but when he assumed the throne of France after the revolution of 1830, Nicholas was consumed with hatred for his former friend who, as he saw it, had gone over to what he perceived as the dark side of liberalism. The idea that Russia was a threat to the East India Company is one version of events. Scholars now favour a different interpretation that the fear of the East India Company was in fact the decision of Dost Mohammed Khan and the Qajar Ruler of Iran to form an alliance and extinguish Sikh rule in Punjab. The British feared that an invading Islamic army would lead to an uprising in India by the people and princely states therefore it was decided to replace Dost Mohammed Khan with a more pliant ruler. "Rumours could cost lives and Empire itself. Therefore instead of fixating on the oriental other, the East India Company played up the threat of the Russian bear".


title: First Anglo-Afghan War

text: The Company sent an envoy to Kabul to form an alliance with Afghanistan's Amir, Dost Mohammad Khan against Russia. Dost Mohammad had recently lost a strategic city of Peshawar to the Sikh Empire and was willing to form an alliance with Britain if they gave support to retake it, but the British were unwilling. Instead, the British feared the "Dal Khalsa", and they considered the Sikh army to be a far more formidable threat than the Afghans who did not have a well-disciplined army, instead of having only a tribal levy where under the banner of "jihad" tribesmen would come out to fight for the Emir. For this reason, Lord Auckland preferred an alliance with the Punjab over an alliance with Afghanistan. The British could have had an alliance with the Punjab or Afghanistan, but not both at the same time.


title: First Anglo-Afghan War

text: When Governor-General of India Lord Auckland heard about the arrival of Russian envoy Count Jan Prosper Witkiewicz (better known by the Russian version of his name as Yan Vitkevich) in Kabul and the possibility that Dost Mohammad might turn to Russia for support, his political advisers exaggerated the threat. Alexander Burnes, the Scotsman who served as the East India Company's chief political officer in Afghanistan, described Witkiewicz: "He was a gentlemanly and agreeable man, of about thirty years of age, spoke French, Turkish and Persian fluently, and wore the uniform of an officer of the Cossacks". The presence of Witkiewicz had thrown Burnes into a state of despair, leading one contemporary to note that he "abandoned himself to despair, bound his head with wet towels and handkerchiefs and took to the smelling bottle". Dost Mohammad had in fact invited Count Witkiewicz to Kabul as a way to frighten the British into making an alliance with him against his archenemy Ranjit Singh, the Maharaja of the Punjab, not because he really wanted an alliance with Russia. The British had the power to compel Singh to return the former Afghan territories he had conquered whereas the Russians did not, which explains why Dost Mohammad Khan wanted an alliance with the British. Burnes wrote home after having dinner with Count Witkiewicz and Dost Mohammad in late December 1837: "We are in a mess home. The emperor of Russia has sent an envoy to Kabul to offer...money [to the Afghans] to fight Rajeet Singh!!! I could not believe my own eyes or ears." On 20 January 1838, Lord Auckland sent an ultimatum to Dost Mohammad telling him: "You must desist from all correspondence with Russia. You must never receive agents from them, or have aught to do with them without our sanction; you must dismiss Captain Viktevitch [Witkiewicz] with courtesy; you must surrender all claims to Peshawar". Burnes himself had complained that Lord Auckland's letter was "so dictatorial and supercilious as to indicate the writer's intention that it should give offense", and tried to avoid delivering it for long as possible. Dost Mohammad was indeed offended by the letter, but in order to avoid a war, he had his special military advisor, the American adventurer Josiah Harlan, engage in talks with Burnes to see if some compromise could be arranged. Burnes in fact had no power to negotiate anything, and Harlan complained that Burnes was just stalling, which led to Dost Mohammad expelling the British diplomatic mission on 26 April 1838.


title: First Anglo-Afghan War

text: British fears of a Persian and Afghan invasion of India took one step closer to becoming a reality when negotiations between the Afghans and Russians broke down in 1838. The Qajar dynasty of Persia, with Russian support, attempted the Siege of Herat. Herat, in Afghanistan, is a city that had historically belonged to Persia; the Qajar shahs had long desired to take it back. It is located in a plain so fertile that is known as the "Granary of Central Asia"; whoever controls Herat and the surrounding countryside also controls the largest source of grain in all of Central Asia. Russia, wanting to increase its presence in Central Asia, had formed an alliance with Qajar Persia, which had territorial disputes with Afghanistan as Herat had been part of the Safavid Persia before 1709. Lord Auckland's plan was to drive away the besiegers and replace Dost Mohammad with Shuja Shah Durrani, who had once ruled Afghanistan and who was willing to ally himself with anyone who might restore him to the Afghan throne. At one point, Shuja had hired an American adventurer - Josiah Harlan - to overthrow Dost Mohammad Khan, despite the fact Harlan's military experience comprised only working as a surgeon with the East India Company's troops in the First Burma War. Shuja Shah had been deposed in 1809 and had been living in exile in British India since 1818, collecting a pension from the East India Company, which believed that he might be useful one day. The British denied that they were invading Afghanistan, claiming they were merely supporting its "legitimate" Shuja government "against foreign interference and factious opposition." Shuja Shah by 1838 was barely remembered by most of his former subjects and those that did viewed him as a cruel, tyrannical ruler who, as the British were soon to learn, had almost no popular support in Afghanistan.


title: First Anglo-Afghan War

text: On 1 October 1838 Lord Auckland issued the Simla Declaration attacking Dost Mohammed Khan for making "an unprovoked attack" on the empire of "our ancient ally, Maharaja Ranjeet Singh", going on to declare that Shuja Shah was "popular throughout Afghanistan" and would enter his former realm "surrounded by his own troops and be supported against foreign interference and factious opposition by the British Army". As the Persians had broken off the siege of Herat and the Emperor Nicholas I of Russia had ordered Count Vitkevich home (he was to commit suicide upon reaching St. Petersburg), the reasons for attempting to put Shuja Shah back on the Afghan throne had vanished. The British historian Sir John William Kaye wrote that the failure of the Persians to take Herat "cut from under the feet of Lord Auckland all ground of justification and rendered the expedition across the Indus at once a folly and a crime". But at this point, Auckland was committed to putting Afghanistan into the British sphere of influence and nothing would stop him from going ahead with the invasion. On 25 November 1838, the two most powerful armies on the Indian subcontinent assembled in a grand review at Ferozepore as Ranjit Singh, the Maharajah of the Punjab brought out the "Dal Khalsa" to march alongside the sepoy troops of the East India Company and the British troops in India with Lord Auckland himself present amid much colorful pageantry and music as men dressed in brightly colored uniforms together with horses and elephants marched in an impressive demonstration of military might. Lord Auckland declared that the "Grand Army of the Indus" would now start the march on Kabul to depose Dost Mohammed and put Shuja Shah back on the Afghan throne, ostensibly because the latter was the rightful Emir, but in reality to place Afghanistan into the British sphere of influence. Speaking in the House of Lords, the Duke of Wellington condemned the invasion, saying that the real difficulties would only begin after the invasion's success, predicting that the Anglo-Indian forces would rout the Afghan tribal levy, only to find themselves struggling to hold on, as the Hindu Kush mountains and Afghanistan had no modern roads, and calling the entire operation "stupid" since Afghanistan was a land of "rocks, sands, deserts, ice and snow".


title: First Anglo-Afghan War

text: British India at this time was a proprietary colony run by the East India Company, which had been granted the right to rule India by the British Crown. India was only one of several proprietary colonies in the British Empire around the world, where various corporations or individuals had been granted the right to rule by the Crown, with for instance Rupert's Land, which was a vast tract covering most of what is now Canada being ruled by the Hudson's Bay Company, but India was easily the most wealthy and profitable of all the proprietary colonies. By the 19th century, the East India Company ruled 90 million Indians and controlled 70 million acres (243,000 square kilometres) of land under its own flag while issuing its own currency, making it into the most powerful corporation in the world. The East India Company had been granted monopolies on trade by the Crown, but it was not owned by the Crown, though the shares in the East India Company were owned by numerous MPs and aristocrats, creating a powerful Company lobby in Parliament while the Company regularly gave "gifts" to influential people in Britain. The East India Company was sufficiently wealthy to maintain the three Presidency armies, known after their presidencies as the Bengal Army, the Bombay Army and the Madras Army, with the supreme field headquarters for commanding these armies being at Simla. The East India Company's army totaled 200,000 men, making it one of the largest armies in the entire world, and was an army larger than those maintained by most European states. The majority of the men serving in the presidency armies were Indian, but the officers were all British, trained at the East India Company's own officer school at the Addiscombe estate outside of London. Furthermore, the politically powerful East India Company had regiments from the British Army sent to India to serve alongside the East India Company's army. Officers from the British Army serving in India tended to look down on officers serving in the company's army, and relations between the two armies were cool at best.


title: First Anglo-Afghan War

text: The regiments chosen for the invasion of Afghanistan came from the Bengal and Bombay armies. The commander in India, Sir Henry Fane, chose the regiments by drawing lots. The units from the Bengal Army going into Afghanistan were Skinner's Horse, the 43rd Native Infantry and the 2nd Light Cavalry, which were all Company regiments while the 16th Lancers and the 13th Somersethire Light Infantry came from the British Army in India. The units from the Bombay Army chosen for the Grand Army of the Indus were the 19th Native Infantry and the Poona Local Horse, which were Company regiments, and the 2nd Queen's Regiment, the 17th Lincolnshire Regiment, and the 4th Dragoons, which were all British Army regiments. Of the two divisions of the Grand Army of the Indus, the Bombay division numbered fifty-six hundred men and the Bengal division numbered ninety-five hundred men. Shuja recruited 6,000 Indian mercenaries ("Shah Shujah's Levy") out of his pocket for the invasion. Ranjit Singh, the elderly and ailing Maharaja of the Punjab and the British assembled in a grand review at Ferozepore as Ranjit Singh, the Maharajah of the Punjab brought out the "Dal Khalsa" to march alongside the sepoy troops of the East India Company and the British troops in India. Ranjit Singh agreed to a treaty with the British viceroy Lord Auckland to restore Shah Shoja to the Afghan throne in Kabul. In pursuance of this agreement, the British army of the Indus entered Afghanistan from the south, while Ranjit Singh's troops went through the Khyber Pass and took part in the victory parade in Kabul. Accompanying the invasion force were 38,000 Indian camp followers and 30,000 camels to carry supplies.


title: First Anglo-Afghan War

text: The Emirate of Afghanistan had no army, and instead under the Afghan feudal system, the tribal chiefs contributed fighting men when the Emir called upon their services. The Afghans were divided into numerous ethnic groups, of which the largest were the Pashtuns, the Tajiks, the Uzbeks, and the Hazaras, who were all in their turn divided into numerous tribes and clans. Islam was the sole unifying factor binding these groups together, though the Hazaras were Shia Muslims while the rest were Sunni Muslims. The Pashtuns were the dominant ethnic group, and it was with the Pashtun tribes that the British interacted the most. The Pashtun tribesmen had no military training, but the ferociously warlike Pashtuns were forever fighting each other, when not being called up for service for the tribal levy by the Emir, meaning most Pashtun men had at least some experience of warfare. The Pashtun tribes lived by their strict moral code of "Pashtunwali" ("the way of the Pashtuns") stating various rules for a Pashtun man to live by, one of which was that a man had to avenge any insult, real or imagined, with violence, in order to be considered a man. The standard Afghan weapon was a matchlock rifle known as the jezail.


title: First Anglo-Afghan War

text: The "Army of the Indus" which included 21,000 British and Indian troops under the command of John Keane, 1st Baron Keane (subsequently replaced by Sir Willoughby Cotton and then by William Elphinstone) set out from Punjab in December 1838. With them was William Hay Macnaghten, the former chief secretary of the Calcutta government, who had been selected as Britain's chief representative to Kabul. It included an immense train of 38,000 camp followers and 30,000 camels, plus a large herd of cattle. The British intended to be comfortable – one regiment took its pack of foxhounds, another took two camels to carry its cigarettes, junior officers were accompanied by up to 40 servants, and one senior officer required 60 camels to carry his personal effects.


title: First Anglo-Afghan War

text: By late March 1839 the British forces had crossed the Bolan Pass, reached the southern Afghan city of Quetta, and begun their march to Kabul. They advanced through rough terrain, across deserts and 4,000-metre-high mountain passes, but made good progress and finally set up camps at Kandahar on 25 April 1839. After reaching Kandahar, Keane decided to wait for the crops to ripen before resuming his march, so it was not until 27 June that the Grand Army of the Indus marched again. Keane left behind his siege engines in Kandahar, which turned out to be a mistake as he discovered that the walls of the Ghazni fortress were far stronger than he expected. A deserter, Abdul Rashed Khan, a nephew of Dost Mohammad Khan, informed the British that one of the gates of the fortress was in bad state of repair and might be blasted open with a gunpowder charge. Before the fortress, the British were attacked by a force of the Ghilji tribesmen fighting under the banner of "jihad" who were desperate to kill "farangis", a pejorative Pashtun term for the British, and were beaten off. The British took fifty prisoners who were brought before Shuja, where one of them stabbed a minister to death with a hidden knife. Shuja had them all beheaded, which led Sir John Kaye, in his official history of the war, to write this act of "wanton barbarity", the "shrill cry" of the "Ghazis", would be remembered as the "funeral wail" of the government's "unholy policy".


title: First Anglo-Afghan War

text: On 23 July 1839, in a surprise attack, the British-led forces captured the fortress of Ghazni, which overlooks a plain leading eastward into the Khyber Pakhtunkhwa. The British troops blew up one city gate and marched into the city in a euphoric mood. During the battle, the British suffered 200 killed and wounded, while the Afghans suffered 500 killed and 1,500 captured. Ghazni was well-supplied, which eased the further advance considerably.


title: First Anglo-Afghan War

text: Following this and an uprising of Tajiks in Istalif, the British marched to Kabul with no resistance from Dost Mohammad's troops. With his situation rapidly deteriorating, Dost Mohammed offered to accept Shuja as his overlord in exchange for becoming his wazir (a common practice in Pashtunwali), which was promptly turned down. In August 1839, after thirty years, Shuja was again enthroned in Kabul. Shuja promptly confirmed his reputation for cruelty by seeking to wreak vengeance on all who had crossed him as he considered his own people to be "dogs" who needed to be taught to obey their master.


title: First Anglo-Afghan War

text: In the absence of the traditional winter capital of Peshawar, on November 2 Shuja left the Bala Hissar to seek refuge from the cold in Jalalabad.


title: First Anglo-Afghan War

text: On 13 November 1839, while en route to India, the Bombay column of the British Indian Army attacked, as a form of reprisal, the Baloch tribal fortress of Kalat, from where Baloch tribes had harassed and attacked British convoys during the move towards the Bolan Pass.


title: First Anglo-Afghan War

text: Dost Mohammad fled to the emir of Bukhara who violated the traditional code of hospitability by throwing Dost Mohammad into his dungeon, where he joined Colonel Charles Stoddart. Stoddart had been sent to Bukhara to sign a treaty of friendship and arrange a subsidy to keep Bukhara in the British sphere of influence, but was sent to the dungeon when Nasrullah Khan decided the British were not offering him a big enough bribe. Unlike Stoddart, Dost Mohammad was able to escape from the dungeon and fled south to Afghanistan. Dost Mohammed fled the dubious hospitality of the embrace of the Emir of Bukhara and on 2 November 1840, his forces turned around at Parwan Darra to meet British general Robert Sale, where he successfully defeated the 2nd Bengal Cavalry. This was principally because the Indians in the 2nd Bengal Cavalry failed to follow their officers who charged towards Dost Mohammed, "The explanation offered by the cavalrymen for not fighting was "that they object to the English sabres" . The simple fact was that despite Britain's industrial revolution, the handcrafted Afghan jezail and sword were far superior to their British counterparts. Indeed, Captain Ponsoby's sword was nearly cut in two."


title: First Anglo-Afghan War

text: Despite Sale having little to show for the campaign and the trail of devastation left by him, Sale called Parwan Darra a victory. However he was unable to conceal the fact of the 2nd Bengal horse defying orders, and as a result, many British officers were killed. Atkinson, the armies surgeon general, called the encounter a “disaster”, Kaye also called the battle a defeat. However, early in the evening of 2 November 1840, a horsemen identified as Sultan Muhammad Khan Safi rode up to Macnaghten, as with this, he was followed by another lone horsemen, who came up to Macnaghten. This horsemen was no other then Dost Mohammad Khan.


title: First Anglo-Afghan War

text: Dost Mohammad's surrender has been given much speculation by historians as to why he surrendered. A number of explanations range from Dost Mohammad's surrender, oddly even after his victory at Parwan Darra. According to Jonathan lee, some of these depictions include Dost Mohammad believing that the British were destined to rule between the Indus and more. Others claim according to Jonathan lee, that, after the defeat at Saighan, Dost Mohammad was free to surrender without loss of honour. Another theory being that Dost Mohammad surrendered due to the brave suicidal charge done by Captain Fraser and Dr Lord, where he could have recognized resistance as futile. This theory however, did not make any sense as where the 2nd Bengal horse faced many defy orders to charge with them, the British lost multiple men due to this and inevitably, the battle. Another theory to Dost Mohammad's surrender was his family being in custody with the British, and about to be exiled to British India. This theory is even less credible, however, due to the fact that the British promising to treat them honourably, as Dost was in no position regardless to change this. Nonetheless, whatever reason Dost Mohammad had surrendered for, it was clear he was not surrendering as a defeated man, as he had successfully defeated the British at Parwan Darra, and even forced their withdrawal from Bamiyan. The revolt in Kohistan also put the military favor within Dost Mohammad's reach, yet for some reason, he decided to give up resistance. Mohan Lal gives the most clear and coherent theory was to why Dost Mohammad surrendered, as according to Lal, Dost Mohammad Khan was plotted against, and in battle, a sniper planned to assassinate him. As further explained by Lal, this was plotted out by Kohistani Amirs, who would then blame the British for this. Lal's accounts and claims are supported by letters to Dost Mohammad Khan from Sultan Muhammad Khan of Nijrab, who had warned the Amir of a plot that was circulating from Maliks who were planning to betray and assassinate him. Abd al-Karim Alawi, who wrote a near contemporary account of the First Anglo-Afghan War, elaborates further on the details of this plot. According to Al-Karim, Sales and Burnes were in contact with the Malik of ‘Ali Hissar, who agreed to kill or capture Dost Mohammad and his two sons in exchange for a substantial payment. According to confidential information shared by British officers, Macnaghten and Burnes had been intercepting communications from Sultan Muhammad Khan of Nijrab, and his supporters. As a result, Macnaghten forged a letter from a well-wisher in Kabul, sending a letter to Dost Mohammad Khan, where it warned the Amir of a plot on his life. Dost Mohammad Khan, having read this letter, was convinced that there was indeed, a plot on his life, as he did not have reason to trust the Kohistanis, as decades prior, he attacked and razed the region. As a result, the only person he seemed to have trusted was Sultan Muhammad Khan of Nijrab. As a result, Dost Mohammad’s options were limited, as the Mir Wali and Murad Beg signed a treaty with Shah Shuja, no longer offering sanctuary for Dost Mohammad. His experience in Bukhara and how he had to escape swayed Dost Mohammad from trying to seek refuge in Balkh, as he could be imprisoned, or even killed. Therefore, he went to the only choice he could think of, the British. The British cantonment at Kabul was only a few hours ride from Parwan Darra, and he had already received assurances that he would be treated honourably in captivity and exile. As a result, Dost Mohammad’s “surrender” is treated by Jonathan Lee and many other historians as nothing akin to surrender, rather seeking refuge with the British government through an old tradition of grabbing the stirrup. An old Turco-Mongolian tradition called rikab giriftan. Atkinson states that Dost Mohammad was seeking the protection of the British government through this action, and as a result, a famous sketch was produced showing Macnaghten and Dost Mohammad shaking hands as a sign of friendship. According to Jonathan Lee, Akbar Khan had later stated that his father “Thrown himself upon the honour of the British government, in times of need”. Macnaghten was elated to hear the news, as according to Jonathan Lee, he stated: “The Afghans are gunpowder and the Dost is a lighted match”. Dost Mohammad was treated honourably, and even was housed in the Bala Hissar, with Dost Mohammad allowed to meet his family, and write to his sons that were in open resistance. Dost Mohammad was also allowed to ride on horse with an escort. During his stay in the Bala Hissar, he held a secret court, with many coming to pay their respects to the former Amir. This had shown Dost Mohammad's popularity, and opposite to Shah Shuja's claims, Shah Shuja was seen as unpopular by many. In Macnaghten's letters, it was shown that he expressed sympathy for Dost Mohammad, showing a very clear contrast to what he referred to him now compared to two years prior, where he denounced Dost Mohammad in the Simla declaration. Macnaghten even asked the Governor General to treat Dost Mohammad better than when they had Shah Shuja in his exile, since to his statement: “We ejected the Dost, who never offended us, in support of our policy, of which he was the victim”.


title: First Anglo-Afghan War

text: The majority of the British troops returned to India, leaving 8,000 in Afghanistan, but it soon became clear that Shuja's rule could only be maintained with the presence of a stronger British force. The Afghans resented the British presence and the rule of Shah Shuja. As the occupation dragged on, the East India Company's first political officer William Hay Macnaghten allowed his soldiers to bring their families to Afghanistan to improve morale; this further infuriated the Afghans, as it appeared the British were setting up a permanent occupation. Macnaghten purchased a mansion in Kabul, where he installed his wife, crystal chandelier, a fine selection of French wines, and hundreds of servants from India, making himself completely at home. Macnaghten, who had once been a judge in a small town in Ulster before deciding he wanted to be much more than a small town judge in Ireland, was known for his arrogant, imperious manner, and was simply called "the Envoy" by both the Afghans and the British. The wife of one British officer, Lady Florentia Sale created an English style garden at her house in Kabul, which was much admired and in August 1841 her daughter Alexadrina was married at her Kabul home to Lieutenant John Sturt of the Royal Engineers. The British officers staged horse races, played cricket and in winter ice skating over the frozen local ponds, which astonished the Afghans who had never seen this before.


title: First Anglo-Afghan War

text: Afghanistan had no army, and instead had a feudal system under which the chiefs would maintain a certain number of armed retainers, principally cavalry together with a number of tribesmen who could be called upon to fight in a time of war; when the Emir went to war, he would call upon his chiefs to bring out their men to fight for him. In 1840, the British strongly pressured Shuja to replace the feudal system with a standing army, which threatened to do away with the power of the chiefs, and which the Emir rejected under the grounds that Afghanistan lacked the financial ability to fund a standing army.


title: First Anglo-Afghan War

text: Dost Mohammad successfully defeated the British at the battle of Parwan Darra on 2 November 1840. However, following his victory, he surrendered and was sent to India in exile after hearing rumours of assassination plots against him.


title: First Anglo-Afghan War

text: In 1839–40, the entire rationale for the occupation of Afghanistan was changed by the Oriental Crisis when Mohammad Ali the Great, the "vali" (governor) of Egypt who was a close French ally, rebelled against the Sublime Porte; during the subsequent crisis, Russia and Britain co-operated against France, and with the improvement in Anglo-Russian relations, the need for a buffer state in Central Asia decreased. The Oriental Crisis of 1840 almost caused an Anglo-French war, which given the long-standing Franco-Russian rivalry caused by Nicholas's detestation of Louis-Philippe as a traitor to the conservative cause, inevitably improved relations between London and St. Petersburg, which ultimately led to the Emperor Nicholas making an imperial visit to London in 1844 to meet Queen Victoria and the Prime Minister Lord Peel. As early as 1838, Count Karl Nesselrode, the Russian Foreign Minister, had suggested to the British Ambassador in St. Petersburg, Lord Clanricarde, that Britain and Russia sign a treaty delimiting spheres of influence in Asia to end the "Great Game" once and for all. By 1840 Clanricarde was reporting to London that he was quite certain a mutually satisfactory agreement could be negotiated, and all he needed was the necessary permission from the Foreign Office to begin talks. From Calcutta, Lord Auckland pressed for acceptance of the Russian offer, writing "I would look forward to a tripartite Treaty of the West under which a limit shall be placed to the advance of England, Russia and Persia and under which all shall continue to repress slave dealing and plunder". Through Britain rejected the Russian offer, after 1840 there was a marked decline in Anglo-Russian rivalry and a "fair working relationship in Asia" had developed. The British Foreign Secretary Lord Palmerston rejected the Russian offer to end the "Great Game" as he believed that as long as the "Great Game" continued, Britain could inconvenience Russia in Asia to better achieve her foreign policy goals in Europe much more than Russia could inconvenience Britain in Asia to achieve her foreign policy goals in Europe. Palmerston noted that because the British had more money to bribe local rulers in Central Asia, this gave them the advantage in this "game", and it was thus better to keep the "Great Game" going. Palmerston believed it was Britain that held the advantage in the "Great Game", that the Russian offer to definitely mark out spheres of influence in Asia was a sign of weakness and he preferred no such treaty be signed. From Palmerston's viewpoint accepting the Russian offer would be unwelcome as the end of the "Great Game" in Asia would mean the redeployment of Russian power to Europe, the place that really counted for him, and it was better to keep the "Great Game" going, albeit at a reduced rate given the tensions with France. At the same time, the lowering of Anglo-Russian tension in the 1840s made holding Afghanistan more of an expensive luxury from the British viewpoint as it no longer seemed quite as essential to have a friendly government in Kabul anymore.


title: First Anglo-Afghan War

text: By this time, the British had vacated the fortress of Bala Hissar and relocated to a cantonment built to the northeast of Kabul. The chosen location was indefensible, being low and swampy with hills on every side. To make matters worse, the cantonment was too large for the number of troops camped in it and had a defensive perimeter almost two miles long. In addition, the stores and supplies were in a separate fort, 300 yards from the main cantonment. The British commander, Major-General George Keith Ephinstone who arrived in April 1841 was bed-ridden most of the time with gout and rheumatism.


title: First Anglo-Afghan War

text: Between April and October 1841, disaffected Afghan tribes were flocking to support resistance against the British in Bamiyan and other areas north of the Hindu Kush mountains. They were organised into an effective resistance by chiefs such as Mir Masjidi Khan and others. In September 1841, Macnaghten reduced the subsidies paid out to Ghilzai tribal chiefs in exchange for accepting Shuja as Emir and to keep the passes open, which immediately led to the Ghazis rebelling and a "jihad" being proclaimed. The monthly subsidies, which were effectively bribes for the Ghazi chiefs to stay loyal, was reduced from 80,000 to 40,000 rupees at a time of rampant inflation, and as the chiefs' loyalty had been entirely financial, the call of "jihad" proved stronger. Macnaghten did not take the threat seriously at first, writing to Henry Rawlinson in Kandahar on 7 October 1841: "The Eastern Ghilzyes are kicking up a row about some deductions which have been made from their pay. The rascals have completely succeeded in cutting communications for the time being, which is very provoking to me at this time; but they will be well trounced for their pains. One down, t'other come on, is the principle of these vagabonds".


title: First Anglo-Afghan War

text: Macnaghten ordered an expedition. On 10 October 1841, the Ghazis in a night raid defeated the Thirty-fifth Native Infantry, but were defeated the next day by the Thirteenth Light Infantry. After their defeat, which led to the rebels fleeing to the mountains, Macnaghten overplayed his hand by demanding that the chiefs who rebelled now send their children to Shuja's court as hostages to prevent another rebellion. As Shuja had a habit of mutilating people who displeased him in the slightest, Macnaghten's demand that the children of the chiefs go to the Emir's court was received with horror, which led the Ghazi chiefs to vow to fight on. Macnaghten, who had just been appointed as the governor of Bombay was torn between a desire to leave Afghanistan on a high note with the country settled and peaceful versus a desire to crush the Ghazis, which led him to temporize, at one moment threatening the harshest reprisals and the next moment, compromising by abandoning his demand for hostages. Macnaghten's alternating policy of confrontation and compromise was perceived as weakness, which encouraged the chiefs around Kabul to start rebelling. Shuja was so unpopular that many of his ministers and the Durrani clan joined the rebellion.


title: First Anglo-Afghan War

text: On the night of 1 November 1841, a group of Afghan chiefs met at the Kabul house of one of their number to plan the uprising, which began in the morning of the next day. In a flammable situation, the spark was provided unintentionally by the East India Company's second political officer, Sir Alexander 'Sekundar' Burnes. A Kashmiri slave girl who belonged to a Pashtun chief Abdullah Khan Achakzai living in Kabul ran away to Burne's house. When Ackakzai sent his retainers to retrieve her, it was discovered that Burnes had taken the slave girl to his bed, and he had one of Azkakzai's men beaten. A secret "jirga" (council) of Pashtun chiefs was held to discuss this violation of "pashtunwali", where Ackakzai holding a Koran in one hand stated: "Now we are justified in throwing this English yoke; they stretch the hand of tyranny to dishonor private citizens great and small: fucking a slave girl isn't worth the ritual bath that follows it: but we have to put a stop right here and now, otherwise these English will ride the donkey of their desires into the field of stupidity, to the point of having all of us arrested and deported to a foreign field". At the end of his speech, all of the chiefs shouted ""Jihad"". November 2, 1841 actually fell on 17 Ramadan which was the anniversary date for the battle of Badr. The Afghans decided to strike on this date for reasons of the blessings associated with this auspicious date of 17 Ramadan. The call to jihad was given on the morning of 2 November from the Pul-i-khisti mosque in Kabul


title: First Anglo-Afghan War

text: Lady Sale wrote in her diary on 2 November 1841: "This morning early, all was in commotion in Kabul. The shops were plundered and the people all fighting." That same day, a mob "thirsting for blood" appeared outside of the house of the East India Company's second political officer, Sir Alexander 'Sekundar' Burnes, where Burnes ordered his sepoy guards not to fire while he stood outside haranguing the mob in Pashto, attempting unconvincingly to persuade the assembled men that he did not bed their daughters and sisters. Captain William Broadfoot who was with Burnes saw the mob march forward, leading him to open fire with another officer writing in his diary that he "killed five or six men with his own hand before he was shot down". The mob smashed in to Burnes's house, where he, his brother Charles, their wives and children, several aides and the sepoys were all torn to pieces. The mob then attacked the home of the paymaster Johnston who was not present, leading to later write when he surveyed the remains of his house that they "gained possession of my treasury by undermining the wall...They murdered the whole of the guard (one officer and 28 sepoys), all my servants (male, female, and children), plundered the treasury...burnt all my office records...and possessed themselves of all my private property". The British forces took no action in response despite being only five minutes away, which encouraged further revolt. The only person who took action that day was Shuja who ordered out one of his regiments from the Bala Hissar commanded by a Scots mercenary named Campbell to crush the riot, but the old city of Kabul with its narrow, twisting streets favored the defenders, with Campbell's men coming under fire from rebels in the houses above. After losing about 200 men killed, Campbell retreated back to the Bala Hissar. After hearing of the defeat of his regiment, Shuja descended into what Kaye called "a pitiable state of dejection and alarm", sinking into a deep state of depression as it finally dawned on him that his people hated him and wanted to see him dead. Captain Sturt was sent to the Bala Hissar by Elphinstone to see if it were possible to recover control of the city later that afternoon, where his mother-in-law Lady Sale noted in her diary: "Just as he entered the precincts of the palace, he was stabbed in three places by a young man well dressed, who escaped into a building close-by, where he was protected by the gates being shut." Sturt was sent home to be cared for by Lady Sale and his wife with the former noting: "He was covered with blood issuing from his mouth and was unable to articulate. He could not lie down, from the blood choking him", only being capable hours later to utter one word: "bet-ter". Lady Sale was highly critical of Elphinstone's leadership, writing: "General Elphinstone vacillates on every point. His own judgement appears to be good, but he is swayed by the last speaker", criticising him for "...a very strange circumstance that troops were not immediately sent into the city to quell the affair in the commencement, but we seem to sit quietly with our hands folded, and look on".." Despite both being in the cantonment, Elphinstone preferred to write letters to Macnaghten, with one letter on 2 November saying "I have been considering what can done tomorrow" (he decided to do nothing that day), stating "our dilemma is a difficult one", and finally concluding "We must see what the morning brings". The British situation soon deteriorated when Afghans stormed the poorly defended supply fort inside Kabul on 9 November.


title: First Anglo-Afghan War

text: In the following weeks, the British commanders tried to negotiate with Akbar Khan. Macnaghten secretly offered to make Akbar Afghanistan's vizier in exchange for allowing the British to stay, while simultaneously disbursing large sums of money to have him assassinated, which was reported to Akbar Khan. A meeting for direct negotiations between Macnaghten and Akbar was held near the cantonment on 23 December, but Macnaghten and the three officers accompanying him were seized and slain by Akbar Khan. Macnaghten's body was dragged through the streets of Kabul and displayed in the bazaar. Elphinstone had partly lost command of his troops already and his authority was badly damaged.


title: First Anglo-Afghan War

text: On 1 January 1842, following some unusual thinking by Major-General William George Keith Elphinstone, which may have had something to do with the poor defensibility of the cantonment, an agreement was reached that provided for the safe exodus of the British garrison and its dependents from Afghanistan. Five days later, the withdrawal began. The departing British contingent numbered around 16,500, of which about 4,500 were military personnel, and over 12,000 were camp followers. Lieutenant Vincent Eyre commented about the camp followers that "These proved from the very first mile a serious clog on our movements". Lady Sale brought with her 40 servants, none of whom she named in her diary while Eyre's son was saved by a female Afghan servant, who rode through an ambush with the boy on her back, but he never gave her name. The American author James M. Perry noted: "Reading the old diaries and journals, it is almost as if these twelve thousand native servants and sepoy wives and children didn't exist individually. In a way, they really didn't. They would die, all of them - shot, stabbed, frozen to death - in these mountain passes, and no one bothered to write down the name of even one of them". The military force consisted mostly of Indian units and one British battalion, 44th Regiment of Foot.


title: First Anglo-Afghan War

text: They were attacked by Ghilzai warriors as they struggled through the snowbound passes. On the first day, the retreating force made only five miles and as Lady Sale wrote about their arrival at a village of Begramee: "There were no tents, save two or three small palls that arrived. Everyone scraped away the snow as best they might, to make a place to lie down. The evening and night were intensely cold; no food for man or beast procurable, except a few handfuls of "bhoosay" [chopped stew], for which we had to pay five to ten rupees". As the night fell and with it, the temperature, dropped well below freezing. The retreating force then learned that they lost all of their supplies of food and their baggage. On the second day all of the men of the Royal Afghan Army's 6th regiment deserted, heading back to Kabul, marking the end of the first attempt to give Afghanistan a national army. For several months afterwards, what had once been Shuja's army was reduced to begging on the streets of Kabul as Akbar had of all of Shuja's mercenaries mutilated before throwing them on the streets to beg. Despite Akbar Khan's promise of safe conduct, the Anglo-Indian force was repeatedly attacked by the Ghilzais, with one especially fierce Afghan attack being beaten off with a spirited bayonet charge by the 44th Foot.


title: First Anglo-Afghan War

text: While trying to cross the Koord-Kabual pass in the Hindu Kush that was described as five miles long and "so narrow and so shut in on either side that the wintry sun rarely penetrates its gloomy recesses", the Anglo-Indian force was ambushed by the Ghilzai tribesmen. Johnson described "murderous fire" that forced the British to abandon all baggage while camp followers regardless of sex and age were cut down with swords. Lady Sale wrote: "Bullets kept whizzing by us" while some of the artillerymen smashed open the regimental store of brandy to get drunk amid the Afghan attacks. Lady Sale wrote she drank a tumbler of sherry "which at any other time would have made me very unlady-like, but now merely warmed me." Lady Sale took a bullet in her wrist while she had to watch as her son-in-law Sturt had "...his horse was shot out from him and before he could rise from the ground he received a severe wound in the abdomen". With his wife and mother-in-law by his side in the snow, Sturt bled to death over the course of the night. The incompetent, naive and gullible Elphinstone continued to believe that Akbar Khan was his "ally", and believed his promise that he would send out the captured supplies if he stopped the retreat on 8 January. Adding to the misery of the British, that night a ferocious blizzard blew in, causing hundreds to freeze to death.


title: First Anglo-Afghan War

text: On 9 January 1842, Akbar sent out a messenger saying he was willing to take all of the British women as hostages, giving his word that they would not be harmed, and said that otherwise his tribesmen would show no mercy and kill all the women and children. One of the British officers sent to negotiate with Akbar heard him say to his tribesmen in Dari (Afghan Farsi) – a language spoken by many British officers – to "spare" the British while saying in Pashto, which most British officers did not speak, to "slay them all". Lady Sale, her pregnant daughter Alexandria and the rest of British women and children accepted Akbar's offer of safe conduct back to Kabul. As the East India Company would not pay a ransom for Indian women and children, Akbar refused to accept them, and so the Indian women and children died with the rest of the force in the Hindu Kush. The camp followers captured by the Afghans were stripped of all their clothing and left to freeze to death in the snow. Lady Sale wrote that as she was taken back to Kabul she noticed: "The road was covered with awful mangled bodies, all naked".


title: First Anglo-Afghan War

text: In the early morning of 10 January, the column resumed its march, with everyone tired, hungry, and cold. Most of the sepoys by this time had lost a finger or two to frostbite, and could not fire their guns. At the narrow pass of Tunghee Tareekee, which was 50 yards long, and only 4 yards wide, the Ghizye tribesmen ambushed the column, killing without mercy all of the camp followers. The Anglo-Indian soldiers fought their way over the corpses of the camp followers with heavy losses to themselves. From a hill, Akbar Khan and his chiefs watched the slaughter while sitting on their horses, being apparently very much amused by the carnage. Captain Shelton and a few soldiers from the 44th regiment held the rear of the column and fought off successive Afghan attacks, despite being outnumbered. Johnson described Shelton as fighting like a "bulldog" with his sword, cutting down any Afghan who tried to take him on so efficiently that by the end of the day no Afghan would challenge him. On the evening of 11 January 1842, General Elphinstone, Captain Shelton, the paymaster Johnston, and Captain Skinner met with Akbar Khan to ask him to stop his attacks on the column. Akbar Khan provided them with warm tea and a fine meal before telling them that they were all now his hostages as he reckoned the East India Company would pay good ransoms for their freedom, and when Captain Skinner tried to resist, he was shot in the face. Command now fell to Brigadier Thomas Anquetil.