Hello! I decided, so to speak, to follow up on my previous article, to write a post where I want to consider in detail the question of why a metronome is needed for a guitarist, and also tell you the structure of a metronome, its main types and purpose.

So, first, we will find out what a metronome is, and then we will move on to the varieties of this device.

Metronome- a mechanical or electronic device that measures (tapping) a certain rhythm at a predetermined speed, ranging from 35 to 250 beats per minute. It is used by musicians when performing a composition as an accurate guide to the tempo and helps during rehearsals when practicing various exercises.

Any piece of music can be played at either a slow or fast tempo. When learning new composition always need to start with slow tempo, in order to ultimately play every note clearly and beautifully. And in this way, gradually approach your goal, reaching the original tempo indicated in the piece of music, thanks to the metronome assistant.

Metronomes are divided into three families:

• Mechanical
• Electronic
• Software

Each musician chooses the metronome that best suits his requirements. Now let's take a closer look at each family.

Mechanical metronomes

The oldest and very first type of metronome that was once invented. The current older generation, who attended music schools in childhood, still remembers small wooden pyramids that stood in glass cabinets or on pianos in the offices of strict music teachers. These pyramids are the ancestors of all modern metronomes.

This species has evolved quite a lot since then. Nowadays, mechanical metronomes are made not only from wood, but also use modern composite materials, such as plastic, for example. Previously, these devices were stationary, but today they are made in a more compact size so that they can easily be put in the pocket of a guitar case.

In the design of some metronomes, special bells began to appear that place emphasis on the downbeat, and this “emphasis” is set depending on the size musical composition, practiced with a metronome. Of course, electronic counterparts are significantly superior in functionality to mechanical metronomes, but the latter have several undeniable advantages that are still worth paying attention to. Here are the main ones:

• Visibility. A mechanical metronome has a pendulum that swings different sides, so it’s hard not to notice it even for a musician who is completely engrossed in playing his instrument. He will always be able to track the movement of the pendulum with peripheral vision.
• Sound. The natural click of a real mechanism cannot be compared with electronics. This sound is absolutely not annoying and can be listened to as a serenade, and it also fits neatly into big picture sound of any instrument.
• Form. For mechanical metronomes it is traditional - in the form of a sophisticated pyramid. This design will add color to any room and also create a creative atmosphere.
• Simplicity. Metronomes of this type, due to their clarity and ease of use, can be used by all musicians without exception, and I would also recommend them to beginning guitarists. They don't need batteries because they have a clock-like mechanism, i.e. Before use, the device must be wound up, like an old mechanical alarm clock.

How does a mechanical metronome work?

The metronome is incredibly simple. The main parts are: steel spring, transmission, anchor escapement. Unlike mechanical watches, the pendulum here is not round, but long with a moving weight, where the axis of the escapement comes into contact with the case and clicks on it. Some models also have a function of strong 2, 3, 5 and 6 beats. Especially for this, the drum is mounted on the release axis, which, like a barrel organ, consists of several wheels with pins, and a bell with a lever moves along it. The bell gives the required share depending on which drum wheel it will be installed opposite.

Electronic metronomes

This is new and modern look metronomes that have captured the hearts of many musicians around the world. Such devices are most preferred by artists who play power instruments. Electronic metronomes, as a rule, are small in size and therefore easily fit in the palm of your hand and can be hidden in any case or travel bag.

Digital metronomes have many useful functions, such as tuning fork, accent, and accent shift, and can satisfy almost any “capricious” user. There are also hybrid models that are combined with a digital tuner, but we will talk about that in another article.

Separately, I would like to mention electronic metronomes for drummers, because... These devices are perhaps the most sophisticated of this family. In addition to various accents and shifts, such metronomes have additional capabilities.

It's no secret that drummers' brains are divided into 4 parts, each of which controls a specific limb. Especially for them, metronomes were invented that can produce a rhythm personally for each limb of the drummer. To do this, the device has several sliders (faders) in order to mix a particular rhythm for a particular leg or arm. This metronome also has a built-in memory for recording and storing rhythms for each individual composition. At concerts, the thing is absolutely irreplaceable - turn on the desired rhythm and tap yourself calmly, being confident that you “can’t run away” from the randomly surging emotions.

From the name it is clear that this is nothing more than special program, installed in the Windows OS environment or an application for Android and iOS. Like real metronomes, virtual metronomes similarly perform their function by generating sound signals at a predetermined pace and/or use visual effects (flashing lights, images of numbers). There are quite a lot of such programs and they are not difficult to find on the Internet.

That's actually all I wanted to tell you in general outline about metronomes. I think now you understand why a guitarist needs a metronome, and you will become friends with it, because... this is very useful and necessary thing in the arsenal of every musician. You will take the right step towards competent guitar playing, because “smooth” musicians have been valued at all times. This is especially appreciated when working together in a group with other musicians. Therefore, I wish you creative heights and success in music. See you soon on the blog pages!

Metronome - now with dance beats!

Don't have a regular metronome? Ours will allow you to learn and rehearse musical pieces in a more comfortable way than with a regular metronome!

If you do not see the metronome above this inscription, then you need to download and install Adobe Flash Player

Good news: Today I received a letter from my childhood friend, classmate, Ivan Lyubchik, with whom I played in a school rock band (Usolye-Sibirskoye, Irkutsk region, 1973-1975). Here's the line: "... Hello Alexey. Yes he uses this metronome all the time … " - Ivan writes about one of his sons - Alexei. Bass guitar player legendary group"Beasts" Alexey Lyubchik rehearses with a Virartek metronome , and Alexey is a very musician high level. So follow the masters!

Online Metronome is very easy to use:

• First button on the left for selection size from the list: 2/4, 3/4, 4/4, 5/4, 7/4, 3/8, 5/8, 6/8, 9/8 and 12/8
• The pace can be set different ways: moving the slider using the " buttons + " And " - ", moving the weight, making several presses in a row on the button " Set the pace"
• volume can be adjusted with a slider
• Can mute the sound and use visual indicators shares: orange- "strong" and blue- "weak"
• you can choose any of 10 sound sets: Wood, Leather, Metal, Time-tick, E-A tones, Tones G-C, Chick-chick, Shaker, Electro, AI Sounds and multiple drum loops for different dance styles, and loops for learning triplets.

To play the drums at the original tempo and size, press the “reset tempo and size” button

Please note that the tempo value is specified for BEATS, i.e. for 4/4 time, 120 would mean 120 quarter notes per minute, and for 3/8 time, 120 eighth notes per minute!

You can “force” the loop to play in a “non-native” time signature, this will give you additional variations in rhythmic patterns.

Sound sets "Tones E-A", "Tones G-C" may be useful for tuning string instrument or for vocal singing.

A large selection of sounds is convenient when using a metronome to learn pieces in different styles. Sometimes you'll need crisp, punchy sounds like AI Sounds, Metal or Electro, sometimes soft sounds like the Shaker set.

A metronome can be useful not only for music lessons. You can use it:

• for learning dance moves;
• doing morning exercises;
• for fast reading training ( certain number blows on time);
• during concentration and meditation.

Musical tempo indications (Wittner metronome scale)
Beats per minute	Italian	Russian
40-60	Largo	Largo - wide, very slow.
60-66	Larghetto	Larghetto is quite slow.
66-76	Adagio	Adagio - slow, calm.
76-108	Andante	Andante - slowly.
108-120	moderato	Moderato - moderate.
120-168	Allegro	Allegro - lively.
168-200	Presto	Presto - fast.
200-208	Prestissimo	Prestissimo - very fast.

Visitor comments:

01.03.2010 Gennady: Correct about the metronome. I would like to know how the tempos that are written in notes (fast, slow, moderate, etc.) relate to the frequency set by the metronome.

01.03.2010 Admin: Especially for you, we have added a sign indicating the tempo of musical works. Please look.

16.05.2010 Irina: Hello! Grandson is 6 years old. He studies music. school. The works are mostly in 2/4 time signature. How to use your metronome in this case. Should the strong beat be on ONE and THREE?

18.05.2010 Admin: Exactly!

02.09.2010 Alexander: Good afternoon, a very high-quality electronic metronome, I’ve been looking for one for a long time. Tell me, is it possible to download it somehow so that I can place it in full screen (without a browser, etc.) and change the background color? I need it for visual use. Thank you.

21.01.2011 Admin: There is no such version yet, but most likely it will appear in February 2011.

23.10.2010 Admin: Almost ALL sizes ADDED!!!

09.11.2010 Valerarv2: Great, that’s all I needed!

13.12.2010 Daria: Guys, I’m in the 7th grade of music. schools. I am preparing for the exams. Thank you very much! All over the World Wide Web I couldn’t find a normal metronome with dimensions! Now I can finally start working out normally :))

20.02.2011 Alex: The long-awaited February is already here. How soon will a computer version of this wonderful metronome appear?

28.02.2011 Svetlana: Super! I love! I would like one like this for my daughter to improve her piano playing. How to purchase this metronome?

03.03.2011 Programmer: A freely available metronome is great. Thank you! And here is the counting " one-and-two-and three-and four-and" would be useful too. Then there are more complex rhythm within, say, the same 4/4 rhythm. The strong part, it seems to me, does not stand out much. It would be nice to do a version with the cymbals hitting the downbeat. Good luck!

05.03.2011 Anton: Thanks for the convenient tool! It's much easier to launch than any professional app just for the sake of a metronome. I often use it for rehearsals and learning parts, and working with students. I would like to ask you to add some sounds (with a sharper attack), as well as loops for training polyrhythms - triplets, doubles, etc. I would also like to have a function for smoothly changing the tempo “FROM and TO”, so that you can practice the part first in a slow, and then at a fast pace...

08.03.2011 Admin: Thank you very much everyone! We really appreciate all suggestions and comments, and we will definitely continue to develop this application. Regarding the desktop version: we are unlikely to release it separately, but Metronome will be included in the package of flash games " Music College"on CD, which is being prepared for release in the near future. Moreover, the applications will work both on Windows and Mac computers.

23.04.2011 Julia: Good day! Thank you very much for the metronome. I’m a teacher at a music school, you won’t find mechanical metronomes during the day, but almost all children have computers. They found you on the Internet. Now a lot of problems have disappeared. All students will become rhythmic))))))))). Thank you, good luck!

In theory, this map should display the places where visitors are located :-)

Anyone who does not play music may consider a metronome to be a useless device, and many do not even know what it is and what its purpose is. The word "metronome" has Greek origin, and it was formed after the merger of two words “law” and “measure”. The invention of the metronome is associated with the name of the great composer Beethoven, who suffered from deafness. The musician relied on the movements of the pendulum to feel the tempo of the piece. The “parent” of the metronome is the Austrian inventor Melzel I.N. Brilliant creator managed to construct a metronome in such a way that it became possible to set the desired tempo of the game.

What is a metronome for?

Metronome- this is a device that plays regular sounds at a certain tempo. By the way, the number of beats per minute can be set independently. Who uses this rhythm machine? For beginners trying to master the guitar, piano, or other instrument, a metronome is a must. After all, while unlearning solo part, you can start the metronome to keep you on track. Music lovers, students music schools and schools, professionals cannot do without a metronome. Even though the metronome sounds like a loud ticking clock, the sound is perfectly audible when playing any instrument. The mechanism counts fractions of a beat and it becomes very convenient to play.

Mechanical or electronic?

Arrived before everyone else mechanical metronomes made from plastic or wood. The pendulum beats the beat, and with the help of the slider a certain tempo is set. The movement of the pendulum is clearly perceptible with peripheral vision. It is worth noting that the main “monsters” musical art prefer mechanical metronomes.

Sometimes they meet metronomes with bell(shown on the left), which emphasizes the downbeat in the measure. The accent can be set according to the size piece of music. The clicks of the mechanical pendulum are not particularly annoying and go well with the sound of any instrument, and anyone can set the metronome.

An undeniable advantage of mechanical devices- independence from batteries. Metronomes are often compared to a clock mechanism: in order for the device to work, it must be wound.

A device with the same functions, but with buttons and a display is electronic metronome. Thanks to its compact size, you can take this device with you on the road. You can find models with a headphone input. This mini metronome can be attached to an instrument or clothing.

Artists playing electronic instruments, choose electrometronomes. The device has a lot of useful functions: accent shift, tuning fork and others. Unlike its mechanical counterpart, the electronic metronome can be set to a “squeak” or “click” if you don’t like the “thump.”

Here is a multifunctional online metronome from the Virartek company, which, among other things, can be used even as a simple drum machine.

How does it work?

A metronome consists of a pendulum with a movable weight and a scale with numbers. If you move the weight along the pendulum, along the scale, the pendulum swings faster or slower and with clicks, similar to the ticking of a clock, marks the desired beat. The higher the weight, the slower the pendulum moves. And if the weight is set in the lowest position, then a quick, as if feverish knock will be heard.

Using the metronome:

Large selection of sizes: click the first button on the left to select a size from the list: 2/4, 3/4, 4/4, etc.
The tempo can be set in different ways: by moving the slider using the “+” and “-” buttons, by moving the weight, by making several presses in a row on the “Set tempo” button
Volume can be adjusted with a slider
You can also turn off the sound and use visual indicators of the beats: orange – “strong” and blue – “weak”
You can choose from 10 sound sets: Wood, Leather, Metal, Raz-tick, E-A Tones, G-C Tones, Chick-Chick, Shaker, Electro, AI Sounds and several drum loops for different dance styles, as well as loops for learning triplets.
To play the drums at the original tempo and size, click the “reset tempo and size” button
The tempo value is indicated for BEATS, i.e. for 4/4 time, 120 would mean 120 quarter notes per minute, and for 3/8 time, 120 eighth notes per minute!
You can force the loop to play in a “non-native” time signature, this will give you additional variations in rhythmic patterns.
Sound sets “Tones E-A”, “Tones G-C” can be useful for tuning a string instrument or for vocal chanting.
A large selection of sounds is convenient when using a metronome to learn pieces in different styles. Sometimes you'll need crisp, punchy sounds like AI Sounds, Metal or Electro, sometimes soft sounds like the Shaker set.

A metronome can be useful for more than just music practice. You can use it:

For learning dance movements;
To train fast reading (a certain number of strokes on a time limit);
During concentration and meditation.

Additional Information:

Musical tempo indications (Wittner metronome scale)

Beats per minute Italian/Russian
40-60 Largo Largo – wide, very slow.
60-66 Larghetto Larghetto is quite slow.
66-76 Adagio Adagio– slowly, calmly.
76-108 Andante Andante - slowly.
108-120 Moderato Moderato – moderate.
120-168 Allegro Allegro is lively.
168-200 Presto Presto – fast.
200-208 Prestissimo Prestissimo – very fast.

How many mechanisms and wonders of technology have been invented by man. And how much he borrowed from nature!.. Sometimes you can’t help but marvel that things from different and seemingly unrelated areas obey general laws. In this article we will draw a parallel between the device that sets the rhythm in music - the metronome - and our heart, which has the physiological property of generating and regulating rhythmic activity.

This work is published as part of a competition for popular science articles held at the Biology - Science of the 21st Century conference in 2015.

Metronome... What kind of thing is this? And this is the same device that musicians use to set the rhythm. The metronome taps beats evenly, allowing you to accurately adhere to the required duration of each measure when performing the entire musical piece. It’s the same with nature: it has had both “music” and “metronomes” for a long time. The first thing that comes to mind when trying to remember what in the body can be similar to a metronome is the heart. A real metronome, isn't it? It also taps beats evenly, even if you play music! But in our cardiac metronome, it is not so much the high accuracy of the intervals between beats that is important, but the ability to constantly maintain the rhythm without stopping. It is this property that will be our main topic today.

So where is the spring responsible for everything hidden in our “metronome”?

Day and night without stopping...

We all know (even more, we can feel) that our heart works constantly and independently. After all, we don’t think at all about controlling the work of the heart muscle. Moreover, even a heart completely isolated from the body will contract rhythmically if nutrients are provided to it (see video). How does this happen? This is an incredible property - cardiac automatism- provided by the conduction system, which generates regular impulses that spread throughout the heart and control the process. That is why the elements of this system are called pacemakers, or pacemakers(from English pacemaker- setting the rhythm). Normally, the heart orchestra is conducted by the main pacemaker - the sinoatrial node. But the question still remains: how do they do it? Let's figure it out.

Contraction of the rabbit heart without external stimuli.

Impulses are electricity. We know where electricity comes from in us - this is the resting membrane potential (RMP) *, which is an indispensable attribute of any living cell on Earth. The difference in ionic composition on different sides of the selectively permeable cell membrane (called electrochemical gradient) determines the ability to generate impulses. Under certain conditions, channels open in the membrane (representing protein molecules with a hole of variable radius), through which ions pass, trying to equalize the concentration on both sides of the membrane. An action potential (AP) arises - the same electrical impulse that propagates along the nerve fibers and ultimately leads to muscle contraction. After the action potential wave has passed, the ion concentration gradients return to their original positions and the resting membrane potential is restored, allowing impulses to be generated again and again. However, the generation of these impulses requires an external stimulus. How then does it happen that pacemakers on one's own generate rhythm?

* - Figuratively and very clearly about the travel of ions through the membrane of a “relaxing” neuron, the intracellular arrest of negative social elements of ions, orphan's share sodium, the proud independence of potassium from sodium and unrequited love cells to potassium, which tends to quietly leak out - see the article “ Formation of the resting membrane potential» . - Ed.

Be patient. Before answering this question, we will have to recall the details of the mechanism for generating an action potential.

Potential - where do opportunities come from?

We have already noted that there is a charge difference between the inner and outer sides of the cell membrane, that is, the membrane polarized(Fig. 1). Actually, this difference is the membrane potential, the usual value of which is about −70 mV (the minus sign means that there is more negative charge inside the cell). The penetration of charged particles through the membrane does not occur by itself; for this, it contains an impressive assortment of special proteins - ion channels. Their classification is based on the type of ions passed through: sodium , potassium , calcium, chlorine and other channels. The channels are capable of opening and closing, but they do this only under the influence of a certain incentive. After stimulation is completed, the channels, like a door on a spring, automatically close.

Figure 1. Membrane polarization. The inner surface of the nerve cell membrane is negatively charged, and the outer surface is positively charged. The image is schematic; details of the membrane structure and ion channels are not shown. Drawing from the site dic.academic.ru.

Figure 2. Propagation of an action potential along a nerve fiber. The depolarization phase is indicated in blue, and the repolarization phase in green. The arrows show the direction of movement of Na + and K + ions. Figure from cogsci.stackexchange.com.

A stimulus is like the doorbell of a welcome guest: it rings, the door opens and the guest enters. The stimulus can be either a mechanical effect or Chemical substance, and electric current (via changes in membrane potential). Accordingly, the channels are mechano-, chemo- and voltage-sensitive. Like doors with a button that only a select few can press.

So, under the influence of a change in membrane potential, certain channels open and allow ions to pass through. This change can vary depending on the charge and direction of movement of the ions. In case positively charged ions enter the cytoplasm, happens depolarization- short-term change in the sign of charges on opposite sides of the membrane (a negative charge is established on the outside, and a positive charge on the inside) (Fig. 2). The prefix “de-” means “movement down”, “decrease”, that is, the polarization of the membrane decreases, and the numerical expression of the negative potential modulo decreases (for example, from the initial −70 mV to −60 mV). When Negative ions enter the cell or positive ions exit, happens hyperpolarization. The prefix “hyper-” means “excess”, and the polarization, on the contrary, becomes more pronounced, and the MPP becomes even more negative (from −70 mV to −80 mV, for example).

But small shifts in the magnetic field are not enough to generate an impulse that will propagate along the nerve fiber. After all, by definition, action potential- This an excitation wave propagating along the membrane of a living cell in the form of a short-term change in the sign of the potential in a small area(Fig. 2). In essence, this is the same depolarization, but on a larger scale and spreading in waves along the nerve fiber. To achieve this effect, use voltage-sensitive ion channels, which are very widely represented in the membranes of excitable cells - neurons and cardiomyocytes. Sodium (Na+) channels are the first to open when an action potential is triggered, allowing these ions to enter the cell along a concentration gradient: after all, there were significantly more of them outside than inside. Those membrane potential values ​​at which depolarizing channels open are called threshold and act as a trigger (Fig. 3).

The potential spreads in the same way: when threshold values ​​are reached, neighboring voltage-sensitive channels open, generating rapid depolarization that spreads further and further along the membrane. If the depolarization was not strong enough and the threshold was not reached, massive channel opening does not occur, and the shift in membrane potential remains a local event (Fig. 3, symbol 4).

The action potential, like any wave, also has a descending phase (Fig. 3, designation 2), which is called repolarization(“re-” means “restoration”) and consists of restoring the original distribution of ions on different sides of the cell membrane. The first event in this process is the opening of potassium (K+) channels. Although potassium ions are also positively charged, their movement is directed outward (Fig. 2, green area), since the equilibrium distribution of these ions is opposite to Na + - there is a lot of potassium inside the cell, and little in the intercellular space*. Thus, the outflow of positive charges from the cell balances the amount of positive charges entering the cell. But in order to completely return the excitable cell to its initial state, the sodium-potassium pump must be activated, transporting sodium outward and potassium inward.

* - To be fair, it is worth clarifying that sodium and potassium are the main ones, but not the only ions, taking part in the formation of the action potential. The process also involves a flow of negatively charged chloride (Cl−) ions, which, like sodium, are more abundant outside the cell. By the way, in plants and fungi, the action potential is largely based on chlorine, and not on cations. - Ed.

Channels, channels and more channels

The tedious explanation of details is over, so let's get back to the topic! So, we have found out the main thing - impulse really does not arise just like that. It is generated by the opening of ion channels in response to a stimulus in the form of depolarization. Moreover, the depolarization must be of such magnitude as to open a sufficient number of channels to shift the membrane potential to threshold values ​​- such that they will trigger the opening of neighboring channels and the generation of a real action potential. But the pacemakers in the heart do without any external stimuli (watch the video at the beginning of the article!). How do they do this?

Figure 3. Changes in membrane potential during different phases of the action potential. MPP is equal to −70 mV. The threshold potential is −55 mV. 1 - ascending phase (depolarization); 2 - descending phase (repolarization); 3 - trace hyperpolarization; 4 - subthreshold potential shifts that did not lead to the generation of a full-fledged impulse. Drawing from Wikipedia.

Remember when we said there was an impressive variety of channels? You really can’t count them: it’s like having separate doors in a house for each guest, and even controlling the entry and exit of visitors depending on the weather and day of the week. So, there are such “doors” that are called low-threshold channels. Continuing the analogy with a guest entering a house, one can imagine that the bell button is located quite high, and in order to ring the bell, you must first stand on the threshold. The higher this button is, the higher the threshold should be. The threshold is the membrane potential, and for each type of ion channel this threshold has its own value (for example, for sodium channels it is −55 mV; see Fig. 3).

So, low-threshold channels (for example, calcium channels) open with very small shifts in the resting membrane potential. To reach the button of these “doors”, you just need to stand on the rug in front of the door. Another interesting property of low-threshold channels: after the act of opening/closing, they cannot open again immediately, but only after some hyperpolarization, which brings them out of the inactive state. And hyperpolarization, except for those cases that we talked about above, also occurs at the end of the action potential, as its last phase (Fig. 3, designation 3), due to the excessive release of K + ions from the cell.

So what do we have? In the presence of low-threshold calcium (Ca 2+ ) channels (LTCs), it becomes easier to generate an impulse (or action potential) after the previous impulse has passed. A slight change in potential - and the channels are already open, letting Ca 2+ cations in and depolarizing the membrane to such a level that channels with a higher threshold are activated and trigger a large-scale development of the AP wave. At the end of this wave, hyperpolarization again puts the inactivated low-threshold channels into a state of readiness.

What if these low-threshold channels did not exist? Hyperpolarization after each AP wave would reduce the excitability of the cell and its ability to generate impulses, because under such conditions, much more positive ions would need to be released into the cytoplasm to achieve the threshold potential. And in the presence of NCC, only a small shift in the membrane potential is enough to trigger the entire sequence of events. Thanks to the activity of low-threshold channels cell excitability increases and the state of “combat readiness” necessary for generating an energetic rhythm is restored faster.

But that's not all. Although the NCC threshold is small, it does exist. So what pushes MPP even to such a low threshold? We found out that pacemakers don’t need any external incentives?! So the heart has it for this funny channels. No, really. That's what they're called - funny channels (from English. funny- “funny”, “amusing” and channels- channels). Why funny? Yes, because most voltage-sensitive channels open during depolarization, and these weirdos open during hyperpolarization (on the contrary, they close during depolarization). These channels belong to a family of proteins that penetrate the membranes of the cells of the heart and central nervous system and have a very serious name - cyclic nucleotide-gated hyperpolarization-activated channels(HCN - hyperpolarization-activated cyclic nucleotide-gated), since the opening of these channels is facilitated by interaction with cAMP (cyclic adenosine monophosphate). Here we have found the missing piece in this puzzle. HCN channels, open at potential values ​​close to the MPP and allowing Na + and K + to pass in, shift this potential to low threshold values. Continuing our analogy, they lay out the missing rug. So the entire cascade of opening/closing channels is repeated, looped and rhythmically self-sustaining (Fig. 4).

Figure 4. Pacemaker action potential. NPK - low-threshold channels, VPK - high-threshold channels. The dashed line is the threshold potential for the military-industrial complex. Different colors The successive stages of an action potential are shown.

So, the conduction system of the heart consists of pacemaker cells (pacemakers), which are capable of autonomously and rhythmically generating impulses by opening and closing a whole set of ion channels. A feature of pacemaker cells is the presence in them of types of ion channels that shift the resting potential to the threshold immediately after the cell reaches the last phase of excitation, which allows for the continuous generation of action potentials.

Thanks to this, the heart also contracts autonomously and rhythmically under the influence of impulses propagating in the myocardium along the “wires” of the conduction system. Moreover, the actual contraction of the heart (systole) occurs during the phase of rapid depolarization and repolarization of pacemakers, and relaxation (diastole) occurs during slow depolarization (Fig. 4). Well, the overall picture of everyone electrical processes in the heart we look at electrocardiogram- ECG (Fig. 5).

Figure 5. Electrocardiogram diagram. Wave P - propagation of excitation through the muscle cells of the atria; QRS complex - propagation of excitation through the muscle cells of the ventricles; ST segment and T wave - repolarization of the ventricular muscle. Drawing from.

Metronome Calibration

It's no secret that, like a metronome, the frequency of which is in the control of the musician, the heart can beat faster or slower. Our autonomic nervous system is such a musician-tuner, and its regulating wheels are adrenalin(towards an increase in contractions) and acetylcholine(towards decreasing). I wonder what changes in heart rate occur mainly due to shortening or prolongation of diastole. And this is logical, because the firing time of the heart muscle itself is quite difficult to speed up; it is much easier to change its resting time. Since diastole corresponds to the phase of slow depolarization, regulation should be carried out by influencing the mechanism of its occurrence (Fig. 6). In fact, this is what happens. As we discussed earlier, slow depolarization is mediated by the activity of low-threshold calcium and "funny" non-selective (sodium-potassium) channels. “Orders” to the vegetative nervous system addressed primarily to these performers.

Figure 6. Slow and fast rhythm of changes in pacemaker cell potentials. With increasing duration of slow depolarization ( A) the rhythm slows down (shown by the dashed line, compare with Fig. 4), while its decrease ( B) leads to an increase in discharges.

Adrenalin, under the influence of which our heart begins to beat like crazy, opens additional calcium and “funny” channels (Fig. 7A). By interacting with β 1 * receptors, adrenaline stimulates the formation of cAMP from ATP ( secondary intermediary), which in turn activates ion channels. As a result, even more positive ions penetrate into the cell, and depolarization develops faster. As a result, the time of slow depolarization is reduced and APs are generated more frequently.

* - The structures and conformational rearrangements of activated G-protein-coupled receptors (including adrenergic receptors), involved in many physiological and pathological processes, are described in the articles: “ A new frontier: the spatial structure of the β 2 -adrenergic receptor has been obtained» , « Receptors in active form» , « β-Adrenergic receptors in active form» . - Ed.

Figure 7. The mechanism of sympathetic (A) and parasympathetic (B) regulation of the activity of ion channels involved in the generation of the action potential of cardiac pacemaker cells. Explanations in the text. Drawing from.

Another type of reaction is observed when interacting acetylcholine with its receptor (also located in the cell membrane). Acetylcholine is an “agent” of the parasympathetic nervous system, which, unlike the sympathetic nervous system, allows us to relax, slow down our heart rate and calmly enjoy life. So, the muscarinic receptor activated by acetylcholine triggers the G-protein conversion reaction, which inhibits the opening of low-threshold calcium channels and stimulates the opening of potassium channels (Fig. 7B). This leads to the fact that fewer positive ions (Ca 2+) enter the cell, and more (K +) exit. All this takes the form of hyperpolarization and slows down the generation of impulses.

It turns out that our pacemakers, although they have autonomy, are not exempt from regulation and adjustment by the body. If necessary, we will mobilize and be fast, and if we don’t need to run anywhere, we will relax.

Breaking is not building

To understand how “dear” certain elements are to the body, scientists have learned to “turn them off.” For example, blocking low-threshold calcium channels immediately leads to noticeable rhythm disturbances: on the ECG recorded on the heart of such experimental animals, there is a noticeable prolongation of the interval between contractions (Fig. 8A), and a decrease in the frequency of pacemaker activity is also observed (Fig. 8B). It is more difficult for pacemakers to shift the membrane potential to threshold values. What if we “turn off” the channels that are activated by hyperpolarization? In this case, mouse embryos will not develop “mature” pacemaker activity (automatism) at all. It’s sad, but such an embryo dies on days 9–11 of its development, as soon as the heart makes its first attempts to contract on its own. It turns out that the described channels play a critical role in the functioning of the heart, and without them, as they say, you can’t go anywhere.

Figure 8. Consequences of blocking low-threshold calcium channels. A- ECG. B- rhythmic activity of pacemaker cells of the atrioventricular node * of a normal mouse heart (WT - wild type) and a mouse of a genetic line lacking the Ca v 3.1 subtype of low-threshold calcium channels. Drawing from.
* - The atrioventricular node controls the conduction of impulses, normally generated by the sinoatrial node, into the ventricles, and with pathology of the sinoatrial node it becomes the main driver of the heart rhythm.

Here is a short story about small screws, springs and weights, which, being elements of one complex mechanism, ensure the coordinated operation of our “metronome” - the pacemaker of the heart. There is only one thing left to do - to applaud Nature for making such a miracle device that serves us faithfully every day and without our efforts!

Literature

1. Ashcroft F. Spark of Life. Electricity in the human body. M.: Alpina Non-fiction, 2015. - 394 pp.;
2. Wikipedia:“Action potential”;Functional roles of Ca v 1.3, Ca v 3.1 and HCN channels in automaticity of mouse atrioventricular cells. Channels. 5 , 251–261;
3. Stieber J., Herrmann S., Feil S., Löster J., Feil R., Biel M. et al. (2003). The hyperpolarization-activated channel HCN4 is required for the generation of pacemaker action potentials in the embryonic heart. Proc. Natl. Acad. Sci. USA. 100 , 15235–15240..