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Saturday, February 14, 2015

Carboniferous rainforest collapse

From Wikipedia, the free encyclopedia

Coal forests continued after the Carboniferous rainforest collapse. These plant fossils are from one of those forests from about 5 million years after the CRC. However, the composition of the forests changed from a lepidodendron dominated forest to one of predominantly tree ferns and seed ferns.

The Carboniferous Rainforest Collapse (CRC) was a minor extinction event that occurred around 305 million years ago in the Carboniferous period.[1] It altered the vast coal forests that covered the equatorial region of Euramerica (Europe and America). This event may have fragmented the forests into isolated 'islands', which in turn caused the extinction of many plant and animal species. After the event, coal-forming tropical forests did continue in large areas of the earth, but their extent and composition were changed.

The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of the Pennsylvanian. The CRC can also be viewed as part of a broader transition of plant species called the "Carboniferous-Permian transition" that continued for another 10 million years into the early Permian. This larger transition has been recognized as one of the two largest extinction events for plant life.[2]

Extinction patterns on land

Ferns and treeferns from Mount Field National Park, giving an impression of how a Carboniferous rainforest might have looked.

In the Carboniferous, the great tropical rainforests of Euramerica supported towering lycopsids, a heterogeneous mix of vegetation, as well as a great diversity of animal life: giant dragonflies, millipedes, cockroaches, amphibians, and the first reptiles.


The rise of rainforests in the Carboniferous greatly altered the landscapes by eroding low-energy, organic-rich anastomosing (braided) river systems with multiple channels and stable alluvial islands. The continuing evolution of tree-like plants increased floodplain stability (less erosion and movement) by the density of floodplain forests, the production of woody debris, and an increase in complexity and diversity of root assemblages.[3]

Collapse occurred through a series of step changes. First there was a gradual rise in the frequency of opportunistic ferns in late Moscovian times.[4] This was followed in the earliest Kasimovian by a major, abrupt extinction of the dominant lycopsids and a change to tree fern dominated ecosystems.[5] This is confirmed by a recent study showing that the presence of meandering and anabranching rivers, occurrences of large woody debris, and records of log jams decrease significantly at the Moscovian-Kasimovian boundary.[3] Rainforests were fragmented forming shrinking islands further and further apart and in latest Kasimovian time, rainforests vanished from the fossil record.


Before the collapse, animal species distribution was very cosmopolitan: the same species existed everywhere across tropical Pangaea, but after the collapse each surviving rainforest 'island' developed its own unique mix of species. Many amphibian species became extinct while reptiles diversified into more species after the initial crisis.[1] These patterns are explained by the theory of island biogeography, a concept that explains how evolution progresses when populations are restricted into isolated pockets. This theory was originally developed for oceanic islands but can be applied equally to any other ecosystem that is fragmented, only existing in small patches, surrounded by another habitat. According to this theory, the initial impact of habitat fragmentation is devastating, with most life dying out quickly from lack of resources. Then, as surviving plants and animals reestablish themselves, they adapt to their restricted environment to take advantage of the new allotment of resources and diversify. After the Carboniferous Rainforest Collapse, each pocket of life evolved in its own way, resulting in a unique species mix which ecologists term endemism.

Extinction patterns in the sea

Stanley and Powell (2003) show a small extinction peak for marine fauna in the Moscovian.[6]

Biotic recovery and evolutionary consequences


The fragmentation of wetlands left a few isolated refugia in Europe; however, even these were unable to maintain the diversity of Moscovian flora.[2] By the Asselian many families that characterized the Moscovian tropical wetlands had disappeared including Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Alethopteridaceae, Cyclopteridaceae, Neurodontopteridaceae.[2]


The depletion of the plant life contributed to the deteriorating levels of oxygen in the atmosphere, which facilitated the enormous arthropods of the time. Due to the decreasing oxygen, these sizes could no longer be accommodated, and thus between this and the loss of habitat, the giant arthropods were wiped out in this event, most notably the giant dragonflies and millipedes (Meganeura and Arthropleura).


Terrestrially adapted early mammal-like reptiles like Archaeothyris were among the groups who quickly recovered after the collapse.

This sudden collapse affected several large groups. Labyrinthodont amphibians were particularly devastated, while the first reptiles fared better, being ecologically adapted to the drier conditions that followed. Amphibians must return to water to lay eggs; in contrast, reptiles – whose amniote eggs have a membrane ensuring gas exchange out of water and can therefore be laid on land – were better adapted to the new conditions. Reptiles acquired new niches at a faster rate than before the collapse and at a much faster rate than amphibians. They acquired new feeding strategies including herbivory and carnivory, previously only having been insectivores and piscivores.[1]

This extinction event had long term implications for the evolution of amphibians. Under prolonged cold conditions, amphibians can survive by decreasing metabolic rates and resorting to overwintering strategies (i.e. spending most of the year inactive in burrows or under logs). However, this is only a short term strategy and not an effective way of dealing with prolonged unfavourable conditions, especially desiccation. Since amphibians had a limited capacity to adapt to the drier conditions that dominated Permian environments, many amphibian families failed to occupy new ecological niches and went extinct.[7]

Possible causes

Climate and Atmosphere

There are several hypotheses about the nature and cause of the Carboniferous Rainforest Collapse, some of which include climate change.[8][9][10] Following a late Bashkirian interval of glaciation, high-frequency shifts in seasonality from humid to arid times began.[11]

Beginning in the latest Middle Pennsylvanian (late Moscovian) a cycle of aridifictaion began. At the time Carboniferous Rainforest Collapse, the climate became cooler and drier, this is reflected in the rock record as the Earth entered into a short, intense ice age. Sea levels dropped by a hundred metres and glacial ice covered most of the southern continent of Gondwana.[12] The cooler, drier climate conditions were not favourable to the growth of rainforests and much of the biodiversity within them. Rainforests shrank into isolated patches, these islands of rainforest were mostly confined to wet valleys further and further apart. Little of the original lycopsid rainforest biome survived this initial climate crisis. The concentration of carbon dioxide in the atmosphere crashed to one of its all time global lows in the Pennsylvanian and early Permian.[12] [11]

Then a succeeding period of global warming reversed the climatic trend; the remaining rainforests, unable to survive the rapidly changing conditions, were finally wiped out.

As the climate aridified again through the later Paleozoic, the rainforests were eventually replaced by seasonally dry biomes.[13] Though the exact speed and nature of the collapse is not clear, it is thought to have occurred relatively quickly in geologic terms, only a few thousand years at most.


After restoring the center of the Skagerrak-Centered Large Igneous Province (SCLIP) using a new reference frame, it has been shown that the Skagerrak plume rose from the core–mantle boundary (CMB) to its ~300 Ma position.[14] The major eruption interval took place in very narrow time interval, of 297 Ma ± 4 Ma. This rift formation coincides with the Moskovian/Kasimovian boundary and the Carboniferous Rainforest Collapse.[15]

Multiple causes

In recent years, scientists have put forth the idea that many of Earth's largest extinction events were due to multiple causes that coincided in time. Proponents of this view suggest multiple causes because they either don't see a single cause as sufficient in strength to cause the mass extinctions or believe that a single cause is likely to produce the taxonomic pattern of the extinction. Two of Earth's largest extinction events have been hypothesized to be multi-causal in nature:

The cause of the Permo-Triassic extinction is unclear and some authors have indicated that it may be best explained by a "Murder on the Orient Express Scenario" where multiple causes contributed to a devastating impact on life. Possible causes supported by strong evidence include the large scale volcanism at the Siberian Traps, the releases of noxious gases, global warming, and anoxia (oxygen depletion) .[16]

David Archibald and David E. Fastovsky discussed a scenario combining three major causes to the K-T extinction: volcanism, marine regression, and extraterrestrial impact, together wiping out the non-avian dinosaurs 66 million years ago.[17] The specific cause of the CRC is not known, but certainly a multiple cause scenario is a possibility.

Timing and Periodicity

The Carboniferous Rainforest Collapse (CRC) was an extinction event that occurred ~307 million years ago, at the end of the Moscovian and the beginning of the Kasimovian stages of the Pennsylvanian.[18] In 1984 Raup and Sepkoski identified a ~26 million year periodicity in the fossil record.[19] Many explanations for this pattern have been proposed including the presence of a hypothetical companion star to the sun,[20] [21] oscillations in the galactic plane, or passage through the Milky Way's spiral arms.[22] The existence of a periodic cycle itself was contentious until 2014 when Melott and Bambach analysed two independent data sets with increased resolution, confirming distinct extinction peaks every 27 million years and specifically, the CRC falls within the maxima of the 27 Myr periodicity cycle (±1.26 Myr).[23]

Climate and geology

Paleosols define a stratigraphic trend that is interpreted to reflect a period of overall decreased hydromorphy, increased free-drainage and landscape stability, and a shift in the overall regional climate to drier conditions in the Upper Pennsylvanian (Missourian), which are consistent with climate interpretations based on contemporaneous paleo-floral assemblages and geological evidence.[24][25]

Fossil sites

Fossil lycopsid, probably Sigillaria, from Joggins, with attached stigmarian roots

Many fossil sites around the world reflect the changing conditions of the Carboniferous Rainforest Collapse.
  • Hamilton, USA
  • Jarrow, UK
  • Linton, USA
  • Newsham, USA
  • Nyrany, Czechoslovakia
  • Joggins, Canada
The Joggins Fossil Cliffs on Nova Scotia's Bay of Fundy, a UNESCO World Heritage Site is a particularly well-preserved fossil site. Fossil skeletons embedded in the crumbling sea cliffs were discovered by Sir Charles Lyell in 1852. In 1859, his colleague William Dawson discovered the oldest known reptile, Hylonomus lyelli, and since then hundreds more skeletons have been found.[26]

Negative feedback

From Wikipedia, the free encyclopedia

A simple negative feedback system descriptive, for example, of some electronic amplifiers. The feedback is negative if the loop gain AB is negative.

Negative feedback occurs when some function of the output of a system, process, or mechanism is fed back in a manner that tends to reduce the fluctuations in the output, whether caused by changes in the input or by other disturbances.

Whereas positive feedback tends to lead to instability via exponential growth or oscillation, negative feedback generally promotes stability. Negative feedback tends to promote a settling to equilibrium, and reduces the effects of perturbations. Negative feedback loops in which just the right amount of correction is applied in the most timely manner can be very stable, accurate, and responsive.

Negative feedback is widely used in mechanical and electronic engineering, but it also occurs naturally within living organisms,[1][2] and can be seen in many other fields from chemistry and economics to physical systems such as the climate. General negative feedback systems are studied in control systems engineering.


Mercury thermostats (circa 1600) using expansion and contraction of columns of mercury in response to temperature changes were used in negative feedback systems to control vents in furnaces, maintaining a steady internal temperature.

In the invisible hand of the market metaphor of economic theory (1776), reactions to price movements provide a feedback mechanism to match supply and demand.

In centrifugal governors, negative feedback is used to maintain a near-constant speed of an engine, irrespective of the load or fuel-supply conditions.

In servomechanisms, the speed or position of an output, as determined by a sensor, is compared to a set value, and any error is reduced by negative feedback to the input.

In audio amplifiers, negative feedback reduces distortion, minimises the effect of manufacturing variations in component parameters, and compensates for changes in characteristics due to temperature change.

In analog computing feedback around operational amplifiers is used to generate mathematical functions such as addition, subtraction, integration, differentiation, logarithm, and antilog functions.

In a phase locked loop feedback is used to maintain a generated alternating waveform in a constant phase to a reference signal. In many implementations the generated waveform is the output, but when used as a demodulator in a FM radio receiver, the error feedback voltage serves as the demodulated output signal. If there is a frequency divider between the generated waveform and the phase comparator, the device acts as a frequency multiplier.

In organisms, feedback enables various measures (eg body temperature, or blood sugar level) to be maintained within a desired range by homeostatic processes.


Negative feedback as a control technique may be seen in the refinements of the water clock introduced by Ktesibios of Alexandria in the 3rd century BCE. Self-regulating mechanisms have existed since antiquity, and were used to maintain a constant level in the reservoirs of water clocks as early as 200 BCE.[3]

The fly-ball governor is an early example of negative feedback.

Negative feedback was implemented in the 17th Century. Cornelius Drebbel had built thermostatically-controlled incubators and ovens in the early 1600s,[4] and centrifugal governors were used to regulate the distance and pressure between millstones in windmills.[5] James Watt patented a form of governor in 1788 to control the speed of his steam engine, and James Clerk Maxwell in 1868 described "component motions" associated with these governors that lead to a decrease in a disturbance or the amplitude of an oscillation.[6]

The term "feedback" was well established by the 1920s, in reference to a means of boosting the gain of an electronic amplifier.[7] Friis and Jensen described this action as "positive feedback" and made passing mention of a contrasting "negative feed-back action" in 1924.[8] Harold Stephen Black came up with the idea of using negative feedback in electronic amplifiers in 1927, submitted a patent application in 1928,[9] and detailed its use in his paper of 1934, where he defined negative feedback as a type of coupling that reduced the gain of the amplifier, in the process greatly increasing its stability and bandwidth.[10][11]

Karl Küpfmüller published papers on a negative-feedback-based automatic gain control system and a feedback system stability criterion in 1928.[12]

Nyquist and Bode built on Black’s work to develop a theory of amplifier stability.[11]

Early researchers in the area of cybernetics subsequently generalized the idea of negative feedback to cover any goal-seeking or purposeful behavior.[13]
All purposeful behavior may be considered to require negative feed-back. If a goal is to be attained, some signals from the goal are necessary at some time to direct the behavior.
Cybernetics pioneer Norbert Wiener helped to formalize the concepts of feedback control, defining feedback in general as "the chain of the transmission and return of information",[14] and negative feedback as the case when:
The information fed back to the control center tends to oppose the departure of the controlled from the controlling quantity...(p97)
While the view of feedback as any "circularity of action" helped to keep the theory simple and consistent, Ashby pointed out that, while it may clash with definitions that require a "materially evident" connection, "the exact definition of feedback is nowhere important".[1] Ashby pointed out the limitations of the concept of "feedback":
The concept of 'feedback', so simple and natural in certain elementary cases, becomes artificial and of little use when the interconnections between the parts become more complex...Such complex systems cannot be treated as an interlaced set of more or less independent feedback circuits, but only as a whole. For understanding the general principles of dynamic systems, therefore, the concept of feedback is inadequate in itself. What is important is that complex systems, richly cross-connected internally, have complex behaviors, and that these behaviors can be goal-seeking in complex patterns. (p54)
To reduce confusion, later authors have suggested alternative terms such as degenerative,[15] self-correcting,[16] balancing,[17] or discrepancy-reducing[18] in place of "negative".


Feedback loops in the human body

In many physical and biological systems, qualitatively different influences can oppose each other. For example, in biochemistry, one set of chemicals drives the system in a given direction, whereas another set of chemicals drives it in an opposing direction. If one or both of these opposing influences are non-linear, equilibrium point(s) result.

In biology, this process (in general, biochemical) is often referred to as homeostasis; whereas in mechanics, the more common term is equilibrium.

In engineering, mathematics and the physical, and biological sciences, common terms for the points around which the system gravitates include: attractors, stable states, eigenstates/eigenfunctions, equilibrium points, and setpoints.

In control theory, negative refers to the sign of the multiplier in mathematical models for feedback. In delta notation, −Δoutput is added to or mixed into the input. In multivariate systems, vectors help to illustrate how several influences can both partially complement and partially oppose each other.[7]

Some authors, in particular with respect to modelling business systems, use negative to refer to the reduction in difference between the desired and actual behavior of a system.[19][20] In a psychology context, on the other hand, negative refers to the valence of the feedback – attractive versus aversive, or praise versus criticism.[21]

In contrast, positive feedback is feedback in which the system responds so as to increase the magnitude of any particular perturbation, resulting in amplification of the original signal instead of stabilization. Any system in which there is positive feedback together with a gain greater than one will result in a runaway situation. Both positive and negative feedback require a feedback loop to operate.

However, negative feedback systems can still be subject to oscillations. This is caused by the slight delays around any loop. Due to these delays the feedback signal of some frequencies can arrive one half cycle later which will have a similar effect to positive feedback and these frequencies can reinforce themselves and grow over time. This problem is often dealt with by attenuating or changing the phase of the problematic frequencies. Unless the system naturally has sufficient damping, many negative feedback systems have low pass filters or dampers fitted.

Some specific implementations

There are a large number of different examples of negative feedback and some are discussed below.

Error-controlled regulation

A regulator R adjusts the input to a system T so the monitored essential variables E are held to set-point values S that result in the desired system output despite disturbances D.[1][22]

One use of feedback is to make a system (say T) self-regulating to minimize the effect of a disturbance (say D). Using a negative feedback loop, a measurement of some variable (for example, a process variable, say E) is subtracted from a required value (the 'set point') to estimate an operational error in system status, which is then used by a regulator (say R) to reduce the gap between the measurement and the required value.[23][24] The regulator modifies the input to the system T according to its interpretation of the error in the status of the system. This error may be introduced by a variety of possible disturbances or 'upsets', some slow and some rapid.[25] The regulation in such systems can range from a simple 'on-off' control to a more complex processing of the error signal.[26]

It may be noted that the physical form of the signals in the system may change from point to point.
So, for example, a change in weather may cause a disturbance to the heat input to a house (as an example of the system T) that is monitored by a thermometer as a change in temperature (as an example of an 'essential variable' E), converted by the thermostat (a 'comparator') into an electrical error in status compared to the 'set point' S, and subsequently used by the regulator (containing a 'controller' that commands gas control valves and an ignitor) ultimately to change the heat provided by a furnace (an 'effector') to counter the initial weather-related disturbance in heat input to the house.

Negative feedback amplifier

The negative feedback amplifier was invented by Harold Stephen Black at Bell Laboratories in 1927, and granted a patent in 1937 (US Patent 2,102,671 "a continuation of application Serial No. 298,155, filed August 8, 1928 ...").[9][27]
"The patent is 52 pages long plus 35 pages of figures. The first 43 pages amount to a small treatise on feedback amplifiers!"[27]
There are many advantages to feedback in amplifiers.[28] In design, the type of feedback and amount of feedback are carefully selected to weigh and optimize these various benefits.

Though negative feedback has many advantages, amplifiers with feedback can oscillate. See the article on step response. They may even exhibit instability. Harry Nyquist of Bell Laboratories proposed the Nyquist stability criterion and the Nyquist plot that identify stable feedback systems, including amplifiers and control systems.

Negative feedback amplifier with external disturbance.[29] The feedback is negative if βA >0.

The figure shows a simplified block diagram of a negative feedback amplifier.
The feedback sets the overall (closed-loop) amplifier gain at a value:
OI=A1+βA1β ,
where the approximate value assumes βA >> 1. This expression shows that a gain greater than one requires β < 1. Because the approximate gain 1/β is independent of the open-loop gain A, the feedback is said to 'desensitize' the closed-loop gain to variations in A (for example, due to manufacturing variations between units, or temperature effects upon components), provided only that the gain A is sufficiently large.[30] In this context, the factor (1+βA) is often called the 'desensitivity factor',[31][32] and in the broader context of feedback effects that include other matters like electrical impedance and bandwidth, the 'improvement factor'.[33]

If the disturbance D is included, the amplifier output becomes:
O=AI1+βA+D1+βA ,
which shows that the feedback reduces the effect of the disturbance by the 'improvement factor' (1+β A). The disturbance D might arise from fluctuations in the amplifier output due to noise and nonlinearity (distortion) within this amplifier, or from other noise sources such as power supplies.[34][35]

The difference signal I–βO at the amplifier input is sometimes called the "error signal".[36] According to the diagram, the error signal is:
Error signal=IβO=I(1βOI)=I1+βAβD1+βA .
From this expression, it can be seen that a large 'improvement factor' (or a large loop gain βA) tends to keep this error signal small.

Although the diagram illustrates the principles of the negative feedback amplifier, modeling a real amplifier as a unilateral forward amplification block and a unilateral feedback block has significant limitations.[37] For methods of analysis that do not make these idealizations, see the article Negative feedback amplifier.

Operational amplifier circuits[edit]

A feedback voltage amplifier using an op amp with finite gain but infinite input impedances and zero output impedance. [38]

Operational amplifier circuits typically employ negative feedback to get a predictable transfer function. Since the open-loop gain of an op-amp is extremely large, a small differential input signal would drive the output of the amplifier to one rail or the other in the absence of negative feedback. A simple example of the use of feedback is the op-amp voltage amplifier shown in the figure.

The idealized model of an operational amplifier assumes that the gain is infinite, the input impedance is infinite, output resistance is zero, and input offset currents and voltages are zero. Such an ideal amplifier draws no current from the resistor divider.[39] Ignoring dynamics (transient effects), the infinite gain of the ideal op-amp means this feedback circuit drives the voltage difference between the two op-amp inputs to zero.[39] Consequently, the voltage gain of the circuit in the diagram, assuming an ideal op amp, is the reciprocal of feedback voltage division ratio β:
A real op-amp has a high but finite gain A at low frequencies, decreasing gradually at higher frequencies. In addition, it exhibits a finite input impedance and a non-zero output impedance.
Although practical op-amps are not ideal, the model of an ideal op-amp often suffices to understand circuit operation at low enough frequencies. As discussed in the previous section, the feedback circuit stabilizes the closed-loop gain and desensitizes the output to fluctuations generated inside the amplifier itself.[40]

Mechanical engineering[edit]

The ballcock or float valve uses negative feedback to control the water level in a cistern.

An example of the use of negative feedback control is the ballcock control of water level (see diagram). In modern engineering, negative feedback loops are found in fuel injection systems and carburettors. Similar control mechanisms are used in heating and cooling systems, such as those involving air conditioners, refrigerators, or freezers.

Biology and chemistry

Control of endocrine hormones by negative feedback.

Some biological systems exhibit negative feedback such as the baroreflex in blood pressure regulation and erythropoiesis. Many biological process (e.g., in the human anatomy) use negative feedback. Examples of this are numerous, from the regulating of body temperature, to the regulating of blood glucose levels. The disruption of feedback loops can lead to undesirable results: in the case of blood glucose levels, if negative feedback fails, the glucose levels in the blood may begin to rise dramatically, thus resulting in diabetes.

For hormone secretion regulated by the negative feedback loop: when gland X releases hormone X, this stimulates target cells to release hormone Y. When there is an excess of hormone Y, gland X "senses" this and inhibits its release of hormone X. As shown in the figure, most endocrine hormones are controlled by a physiologic negative feedback inhibition loop, such as the glucocorticoids secreted by the adrenal cortex. The hypothalamus secretes corticotropin-releasing hormone (CRH), which directs the anterior pituitary gland to secrete adrenocorticotropic hormone (ACTH). In turn, ACTH directs the adrenal cortex to secrete glucocorticoids, such as cortisol. Glucocorticoids not only perform their respective functions throughout the body but also negatively affect the release of further stimulating secretions of both the hypothalamus and the pituitary gland, effectively reducing the output of glucocorticoids once a sufficient amount has been released.[41]


Self-organization is the capability of certain systems "of organizing their own behavior or structure".[42] There are many possible factors contributing to this capacity, and most often positive feedback is identified as a possible contributor. However, negative feedback also can play a role.[43]


In economics, automatic stabilisers are government programs that are intended to work as negative feedback to dampen fluctuations in real GDP.

Free market economic theorists claim that the pricing mechanism operated to match supply and demand. However Norbert Wiener wrote in 1948:
"There is a belief current in many countries and elevated to the rank of an official article of faith in the United States that free competition is itself a homeostatic process... Unfortunately the evidence, such as it is, is against this simple-minded theory."[44]