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Витамины: строение, свойства, использование, значение для растений

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27

Ужгородський національний університет

Біологічний факультет

Кафедра генетики і фізіології рослин

Біланич М.М.

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27

Ужгород, 2007

ЗМІСТ

ВСТУП………………………………………………………….
…………………………………………3

1. Характеристика вітамінів: будова, властивості,
поширення…………….4

2. Фізіологічна роль вітамінів у життєдіяльності рослин……………….14

ЛІТЕРАТУРА……………………………………………………..
……………………………………25

Вступ

Вітаміни (від лат. Vita- життя) – низькомолекулярні органічні сполуки
різної хімічної природи, які виконують важливі біохімічні і фізіологічні
функції в живих організмах. Для організму потрібні в дуже невеликих
кількостях (від декількох мікрограм до кількох міліграм на добу), тому
що володіють високою біологічною активністю.

Потрібні вітаміни всім живим організмам. Є організми, які здатні
виробляти вітаміни, а також такі, що не можуть їх синтезувати, або не
можуть синтезувати окремі вітаміни. Це, в основному, деякі тварини. В їх
числі людина, пацюки та інші. Вітаміни, звичайно, необхідні і рослинам.
Вони є продуцентами цих речовин. Але Овчаров К.Є. встановив, що є
можливість виникнення авітамінозного і гіповітамінозного стану у рослин.
Ним встановлено, що ці аномальні стани у рослин супроводжуються
серйозними біохімічними порушеннями: подавляється активність
аскорбіноксидази, поліфенолоксидази і пероксидази, затримується
біосинтез вітамінів понижується поступання фосфору і азоту з поживного
середовища (Строганов, Генкель,1976).

Важливим є те, що в продуцентах вітаміни можуть бути в формі
провітамінів. Коли тварини споживають рослини, то провітаміни в
тваринному організмі перетворюються у вітаміни.

Особливо вітаміни необхідні для насіння, яке проростає, що з тих чи
інших причин може витратити передчасно запас вітамінів, який у ньому
відклався при формуванні.

Також важливими продуцентами вітамінів (для людини) є мікроорганізми.
По-перше деякі з них являються симбіотичними з нашим організмами, які
живуть в кишечнику і постачають нас деякими вітамінами. По-друге, часто
людина використовує мікроорганізми для синтезу цих біоактивних речовин в
фармацевтиці, сільському господарстві (для тваринних кормів), у наукових
дослідженнях (біотехнології) та з іншими цілями.

Розглянемо, які є вітаміни, їхні будову, біохімічні і фізіологічні
властивості.

1. Характеристика вітамінів: будова, властивості, поширення

Жиророзчинні вітаміни.

1) Каротини. Вітамін А (Ретинол)

В рослинних продуктах вітамін А у
вигляді ретинолу не накопичується. Рослини синтезують лише
каротиноїди – провітаміни А. Каротиноїди – це пігменти, що
надають овочам і фруктам жовте
забарвлення або червоне. Із всіх каротинів
найбільшою активністю володіє в-каротин, що
перевищує активність Ь- і г- каротинів в 2-3 рази
(Дунаєвський, 1990).

В організмах тварин в-каротин розщеплюється з
утворенням двох молекул ретинола (Кретович, 1986).

СН3 СН3 СН3 СН3

СН = СН – С = СН – СН = СН – С = СН – СН2ОН

СН3

Ретинол.

Каротин термостійкий тому добре зберігається при тепловій обробці
продуктів. Але каротин в 3 рази слабкіший ніж ретинол. Добова потреба
людини в вітаміні А – 1,5-2,5 мг. Забезпечується добова потреба на 2/3
овочами і фруктами. З фруктів і ягід на каротин багаті абрикос і
обліпиха, а менш багаті – вишні, сливи, горобина чорноплідна, малина,
червона смородина та інші.

2) Вітамін К. Цей вітамін міститься в основному в рослинних
продуктах, причому у більш активній формі, ніж у продуктах тваринного
походження (Дунаєвський, 1990).

По всій хімічній природі вітаміни групи К являють собою похідні
нафтохінона (Кретович, 1986).

Вітамін К1, що міститься в зелених листях (хлоропластах), має
таку структуру:

0

СН3 СН3 СН3 СН3

Н2 – СН = С – (СН2)3 – СН – (СН2)3 – С

0 СН3

Вітамін К1.

У організмах людини і тварин грає важливу роль у процесах
згортання крові. Він стимулює в людському організмі синтез протромбіну,
фібриногену, утворення тромбопластину й тромбіну, показує позитивний
вплив на тонус судин. Добова потреба в (філохіноні) вітаміні К для
дорослої людини сягає приблизно 0,2-0,3мл.

Джерелом вітамінів з фруктів і ягід є суниці, яблука, виноград
(Дунаєвський, 1990).

3) Вітаміни групи Е. Токофероли. В овочах і фруктах знаходяться в
порівняно невеликих кількостях: 0,1-0,6 мг/100г (Дунаєвський, 1990).

Хімічна формула:

СН3 О СН3 СН3 СН3 СН3

НО С – (СН2)3 – СН – (СН2)3 – СН – (СН2)3 – СН

3HС СН3

СН3

Вітамін Е.

Цей вітамін бере участь у всіх видах обміну, має властивість
ліотропної дії, стимулює діяльність серцевого м’язу у людини і тварин.
Хоча механізм дії вітаміну Е на молекулярному рівні ще повністю не
розшифровано, тим не менше багато проявів недостачі цьго вітаміну
можуть бути зв`язані з пошкодженнями мембран і клітинних органел, які
при цьому виникають ( Спиричев, Конь,1978).

4) Провітамін Д – стероли.

Вітаміни групи Д зустрічаються лише в тваринних організмах, В
рослинах містяться стероли, з яких під дією ультрафіолетового
опромінення утворюються вітаміни цієї групи. Найбільш важливим з цих
стеролів являється ергостерол, що в великій кількості міститься в
дріжджах і плісневих грибах.

Із ергостерола при його ультрафіолетовому опроміненні утворюється
вітамін Д2 (Кретович, 1986).

Водорозчинні вітаміни.

1) Вітамін С. Аскорбінова кислота.

О О

С С

ОН – С – 2 Н О = С

ОН – С О О = С О

Н – С + 2 Н Н – С

ОН – С – Н ОН – С – Н

СН2ОН СН2ОН

відновлена форма окислена форма

(L-дегідроаскорбінова кислота)

Аскорбінову кислоту можна зобразити ще так:

СН2 – ОН

Н ОН

С

ОН

С – С

О

С – С

ОН О

Хімічні формули вітаміну С (Богатих, 1989)

Виділений угорським вченим Сандьорді в 1936 році. Для людини
добова норма 50-100 мг. Міститься в шипшині, цитрусових, чорній
смородині, у великих кількостях також міститься в обліписі і в сливах, у
меншій кількості у черешнях, малині, винограді, в грецьких горіхах. В
організмі деяких тварин і людини впливає на синтез колагена.

Важлива роль аскорбінової кислоти і її участь в окисно-відновних
процесах, що проходять в живій клітині, зв’язані з тим, що вітамін існує
в двох формах – власне аскорбінової кислоти і легко утворюючоїся з неї
при окисленні дегідроаскорбінової кислоти; остання при відновленні знову
дає аскорбінову кислоту (Кретович, 1986).

У клітинах також є аскорбіноген, який являється зв’язаною формою
аскорбінової кислоти. Ця сполука найбільш стійка до окислення, однак
вітамінна активність його в 2 рази слабкіша за активність аскорбінової
кислоти. Дегідроаскорбінова кислота нестійка й при подальшому її
окисленні втрачає вітамінні властивості.

Потреба людини в аскорбіновій кислоті в основному
задовольняється за рахунок картоплі і капусти, а фрукти і ягоди служать
лише допоміжним джерелом.

В покривних тканинах фруктів вміст аскорбінової кислоти вищий.
Так в кожурі мандаринів, лимонів, апельсинів, яблук рівень її рівний
відповідно 130, 140, 170 і 40 мг/100г, а в плодовій м’якоті мандаринів –
38, лимонів – 66, апельсинів – 55, яблук – 12-13мг/100г. Коливання
кількості аскорбінової кислоти в різних сортах фруктів і овочів більше,
ніж інших вітамінів (Дунаєвський, 1990).

Аскорбінова кислота являє собою безкольорові кристалики кислого
смаку, добре розчиняється в воді, легко руйнується в розчинах, особливо
при присутності повітря, світла і слідів міді або заліза (Кретович,
1986). Також можуть бути значні втрати вітаміну при зберіганні овочів і
фруктів. В овочах і фруктах аскорбінова кислота має сильнішу біологічну
дію, ніж в тваринних продуктах бо знаходиться в контакті з
біофлавоноїдами, що посилюють дію цього вітаміну і застерігають його від
пошкоджень.

2) Вітамін В1

Н ОН

N C–СН2 – N

СН3 N НС С – СН2 – СН2ОН

S

Вітамін В1.

Відкрив Казимір Фунг. Входить вітамін В1 до складу коферментів,
наприклад до ТДФ – тіаміндифосфат або тіамінпірофосфат, ферментів ліаз.
У плодах і овочах міститься десь 0,02 – 0,1мг/100г, а в фруктах трохи
менше: 0,01 – 0,06мг/100г (Дунаєвський, 1990). Грає роль в
вуглеводневому обміні тварин, рослин і бактерій, так як входить в склад
фермента піруватдекарбоксилази, що розчеплює піровиноградну кислоту, яка
утворюється при дисиміляції вуглеводів.

В кислому середовищі тіамін стійкий до нагрівання і кип’ятіння,
але легко руйнується при нагріванні в нейтральному і лужному середовищі
(Кретович, 1986)

3) Вітамін В2. Рибофлавін. Добова норма людини до 2 міліграм. У
будову вітаміну входить ізоалоксазинове кільце:

СН2 – (СНOH)3 – СН2ОН

N N O

СН3

СН3 N

Вітамін В2.

Входить до складу аеробних дегідрогеназ: ФАД, ФМН, як кофермент.
Авітаміноз у людини викликає катаракту. Гіповітаміноз викликає порушення
апетиту, слабкість, больові відчуття в слизових рота і ріжучі відчуття в
очах. (Кретович, 1986).

У плодоягідних культурах накопичується в малих кількостях.
Наприклад, у фруктах в середньому 0,01-0,06 мг/100гр (Дунаєвський,
1990).

4) Вітамін Вс. Фолієва кислота

–––

–––

ОН

N СН2 – N – – C

H

NН2 N N

Фолієва кислота.

Вітамін В6 включає декілька біоактивних речовин:
параамінбензойну кислоту, глутамінову кислоту, які входять в хімічну
будову вітаміну. Вперше фолієва кислота була виділена з лисків шпинату
(лат. follium – листок).

В рослинах вітамін знаходиться переважно в зв’язаній формі й
немає у цій формі біологічної активності. Потрапляючи з харчами у
дванадцятипалу кишку, переходить в вільну форму під дією ферментів.
Вітамін входить в склад ферментів, що беруть участь в біосинтезі
метіоніна, пуринових основ. Рослинними джерелами фолієвої кислоти
являються листові овочі, картопля, яблука, абрикоси. Містяться і
грецьких горіхах. В перестиглих плодах рівень її вищий. (Дунаєвський,
1990). Добова потреба в фолієвій кислоті у людини становить 200-300 мкг.
Фолієва кислота позитивно впливає на лікування деяких важких форм анемії
і малокрів’я людини. Похідні фолієвої кислоти, сама фолієва кислота і
споріднені їй сполуки називають фолатами (Богатих, 1989).

5) Вітамін В3. Пантотенова кислота

Н СН3 ОН О

НО – С – С – С – С – N – СН2 – СН2 – СООН

Н СН3 Н

Вітамін В3. 2-метил – 3 – оксімасляна кислота.

Дуже поширена в продуктах, тому і має таку назву (лат. Panteas –
всюдисущий). У вигляді коферменту А, вона входить в склад багатьох
ферментів, і являється складовою частиною ацетилкоензиму А, який у
людському організмі бере участь у синтезі багатьох речовин: холестерину,
ацетилхоліну, гемоглобіну, а також являється важливим джерелом енергії.
Цей вітамін функціонально пов’язаний з фолієвою кислотою і біотином.

Добова норма пантотенової кислоти для людини становить 5-10мг. З
фруктів і ягід нею багаті чорна смородина, гранати, абрикоси, грецькі
горіхи, персики та інше. Вітамін не стійкий, бо при тепловій обробці
втрати його становлять 20-40%.

5) Вітамін В6. Пірідоксин.

С – Н

СН2 – ОН

Н3С N

Пірідоксин.

Пірідоксин у вигляді коферменту входить в склад ферментів, які
регулюють білковий обмін. Необхідний для нормальної діяльності ЦНС
тварин і людини. Потреба здорової людини в пірідоксині 2-3мг на добу. У
багатьох овочах і фруктах, у яких він міститься, рівень його сягає
0,1-0,2мг/100гр. В дещо більшій кількості цього вітаміну можуть бути в
гранатах, бананах і горіхах. Вітамін добре зберігається при тепловій
обробці (Дунаєвський, 1990). Відсутність чи недостача вітаміну В6 в
харчах приводить до порушень білкового обміну і синтезу жирів в
тваринному організмі (Кретович, 1986).

6) Вітамін В7.(Н). Біотин

N

O=C S

N C?CH2?CH2?CH2?COOH

H H

Вітамін Н.

Вітамін стійкий до нагрівання, окислення, дії кислот і основ. В
фруктах міститься в невеликих кількостях: 0,1-2,0мг/100гр. Трохи більше
за середню концентрацію його є в чорній смородині, винограді,
апельсинах. Добова норма його для людини 0,2-0,3мг на добу в зв’язку з
високою активністю.

7) Нікотинова кислота. Ніацин. Вітамін РР або В5.

С

N

Хімічна формула нікотинової кислоти.

Якщо в даній формулі групу -СООН замістити на групу -NН2, то
утворюються формула аміду нікотинової кислоти. Термін амід нікотинової
кислоти потрібно називати нікотинамідом (Богатих, 1989).

Широко поширений цей вітамін в продуктах рослинного і тваринного
походження. З плодів і ягід багаті ним абрикоси, персики, вишні, малина
та деякі інші.

Ніацин входить в склад окисно-відновних ферментів, бере участь у
вуглеводневому й білковому обмінах. Нікотинова кислота добре
зберігається в овочах і фруктах при тепловій обробці їх, сушці і
консервуванні (Дунаєвський,1990).

Вітаміноподібні речовини.

До вітаміноподібних речовин відносяться ряд сполук, що входять в склад
рослинних продуктів, які по своїй ролі в організмі наближаються до
вітамінів.

1) Параамінбензойна кислота. Вітамін В10

NH2 COOH

Хімічна формула ПАБК.

Параамінбензойна кислота являється складовою фолієвої кислоти
(Вс), разом з якою утворює коферменти необхідні для синтезу нуклеїнових
кислот, метіоніна, інших біологічно активних речовин. Вона стійка при
нагріванні і зберігає активність при довгій тепловій обробці продуктів.
(Дунаєвський, 1990).

В рослинах і тваринних тканинах пара-амінбензойна кислота
головним чином зв’язана з білками, поліпептидами та амінокислотами, а
також міститься в вигляді ацетильного похідного (Кретович, 1986).

2) Вітамін Р. Біофлавоноїди.

Біофлавоноїди відносяться до поліфенолів – складові органічних
сполук, що мають в основі ароматичне кільце з двома і більше
гідроксильними групами. Широко поширені в рослинному світі, біологічно
високоактивні, грають важливу у житті рослин. В наш час число відомих
поліфенолів сягає 2000 і більше.

З багатьох поліфенолів важливе значення мають флавоноїди. Основу
їх структури складає флавон.

В рослинах флавоноїди існують у вільному стані і у стані, коли
зв’язані з цукрами – глікозиди.

Вони регулюють процес росту рослин і грають роль в процесах
біологічного окислення. Флавоноїди поділяються на декілька груп в
залежності від хімічної структури. Частіше вони розчинені в клітинному
соці, рідше знаходяться в клітинних оболонках. В тваринному організмі
вони володіють Р-вітамінною активністю і надають укріплюючу дію на
стінки капіляр і знижують ламкість судин. Вперше були виділені з
червоного перцю у 1936 році.

До біофлавоноїдів відносяться антоціани. Кожен антоціан має своє
забарвлення і в залежності від рН клітинного соку має свій відтінок.
Наявність великої кількості різних антоціанів надає плодам і квітам
різного забарвлення (чорна і червона смородина, малина, суниця,
виноград, абрикоси і інші) (Дунаєвський, 1990).

До речовин, що мають Р-вітамінну активність, відносять широко
поширені в рослинному світі глікозиди – рутин і геспередин, а також
танін, що міститься в чайному листі і винограді. (Кретович, 1986).

3) Ліпоєва кислота. Протоген. Вона являє собою циклічний дисульфід, що
має наступну будову.

CH2 O

H2C CH – CH2 – CH2 – CH2 – CH2 – C

OH

S S

Ліпоєва кислота.

Широко розповсюджена в природі й міститься в більшості продуктів
харчування. Рівень ліпоєвої кислоти вищий у зеленій частині рослин. Є
гіпотези й теорії про важливість участі ліпоєвої кислоти в процесі
фотосинтезу. (Кретович, 1986).

4) Вітамін U. S-метилметіонін.

В плодах і ягодах міститься в невеликих кількостях, а в овочах у
трохи більших кількостях. Наприклад, в помідорах 11-29мг/100гр. Вітамін
термолабільний в зв’язку з чим є його значні втрати при тепловій обробці
продуктів. (Дунаєвський, 1990).

2. Фізіологічна роль вітамінів у життєдіяльності рослин

Накопичені за останні десятиріччя факти показують, що вітаміни так само
потрібні рослинам як і тваринам. Це не просто побічні продукти їх обміну
речовин, а фізіологічно активні речовини, що беруть участь в
життєво-важливих процесах. Далі вияснили, що рослини теж можуть мати
вітамінну недостатність, хоча вона й не дуже значна, але якщо добавляти
вітаміни в грунт або проводити вітамінні підкормки, то це дозволяє
помітно підвищувати продуктивність рослин. (Овчаров, 1962).

Доказом того, що рослинам потрібні вітаміни може бути й такий
факт, що при обробці насіння антивітамінами деяких вітамінів,
відбувається затримка проростання або ж і зменшення схожості насіння.
Також може відбуватися пригнічення росту органів проростків і, взагалі,
загибель проростків. За даними деяких авторів (Смашевський, Копелевич,
Гунар, 1986), Д-гомопантотенова кислота, яка являється антивітаміном
пантотенової кислоти, в концентрації 1 г/л понижувала схожість насіння
огірка на 24,5%, томату –24,1%, рису – 43,8% і кукурудзи – 5,3%. Також у
цій концентрації цей антивітамін особливо пригнічував ріст проростків
цих рослин. Пантотенова кислота у молярному відношенні 1 : 2 повністю
знімала антивітамінну дію гомопантотенової кислоти.

Експерименти показують, що за допомогою вітамінів можна
керувати деякими процесами: у рослин – посилювати або затримувати їх
ріст, прискорювати утворення плодів і т.д.

Як відомо у всіх живих організмах є біологічні
каталізатори-ферменти, дуже малі кількості яких в сотні тисяч разів
прискорюють величезні кількості біохімічних реакцій.

Тепер твердо встановлено, що всі ферменти – білкові речовини.
Вони складаються з білкового “носія” і особливої активної групи –
коферменту, який вступає в хімічну взаємодію з субстратом.

Цією активною групою багатьох ферментів є вітаміни. В залежності
від того з якими білками сполучається кофермент – вітамін, утворюється
той чи інший фермент. Так, наприклад, вітаміни В2 здатен сполучатися
більш, ніж з 20-ма білками (сполучається), утворюючи відповідне число
ферментів з різними фізіологічними функціями. Вітамін В1 разом з двома
молекулами фосфорної кислоти утворює активну групу карбоксилази –
ферменту, дуже широко поширеного в рослинах і організмах тварин,
необхідного для перетворення вуглеводів.

Вітаміни беруть участь в різноманітних процесах перетворення
речовин і життєздатності рослинного організму.

У процесі фотосинтезу, на всіх етапах його, ми зустрічаємося з
вітамінами. В першу чергу це провітаміни А – каротини: вони завжди є
супутниками хлорофілу, бо зажди знаходяться в хлорофілових зернах. Зараз
відомо, що каротин наряду з хлорофілом бере участь в поглинанні енергії
світла. Крім того він береже хлорофіл від розкладання. Вітамін С також
захищає зелений пігмент від окислення. Крім того цей вітамін С бере
участь разом з вітаміном К у складних синтетичних реакціях, що проходять
при фотосинтезі. Також у процесі фотосинтезу беруть участь вітаміни: В6,
РР, біотин та інші. В подальшому перетворенні продуктів фотосинтезу бере
участь вітамін В1. В ці процеси входить також процес під назвою “темнова
фіксація” вуглекислого газу. Цей процес по результатам аналогічний
фотосинтезу і проходить без світла за рахунок енергії хімічних сполук.
Без вітаміну В1 він порушується.

Значну роль виконують вітаміни в перетворенні фосфору. Сполуки
фосфору в усіх живих системах служать акумуляторами енергії. Накопичена
в реакційноздатних фосфорних сполуках енергія після цього
використовується в інших реакціях. Ось у ці фосфорні сполуки входять
багато вітамінів. Їх присутність важлива для зв’язування великої
кількості фосфорної кислоти, що поступила в рослину. Таку роль виконує,
наприклад, інозит. Деякі експерименти показали, що вітамін РР, якщо його
внести в грунт, теж збільшує поглинання рослинами фосфору (Овчаров,
1962).

Вітамін В1 бере участь у перетвореннях сірки. Як відомо в
корінні рослин проходять перетворення сульфатів, що поступають з
грунтового розчину, в вітамін В1 і в амінокислоти.

Величезна роль вітамінів в диханні. Дихання рослин, як і у
тварин, являється найважливішим джерелом енергії для всіх процесів:
синтезу, росту, руху і т.д. Разом з тим при диханні утворюються важливі
для організму сполуки. Дихання проходить в організмі за допомогою
складної системи ферментів.

Також вітаміни грають важливу роль у стійкості рослин. Наприклад
встановлено, що вітаміни групи В – В1, В2, В6 і Вс в концентраціях 1,9 ·
10?іМ, 2,6 · 10О4 М, 2,9 · 10ОіМ, 2,8 · 10Оі М, відповідно, знімають
пригнічуючу дію гербіциду 2,4 – Д на ріст і розвиток рослин, сприяють
мобілізації і посиленню метаболічних процесів відновлення
ультраструктури рослинних клітин при дії цього гербіциду, а також грають
роль у виведенні деякої її частини за допомогою механізму екзоцитозу
(Буадзе,1987)

В склад ферментів дихання входить вітамін В2, часто він
зв’язаний з фосфорною кислотою (Н3 PO4).

Велику роль в процесах дихання грають і вітаміни С, В1, РР,
фолієва кислота, біотин, пантотенова кислота та інші. Кожний вітамін, що
показаний вище, входить в склад відповідного ферменту і виконує властиву
йому функцію в складному процесі, яким являється дихання. Тому не
випадково при проростанні насіння має місце енергійне утворення
вітамінів. Адже енергія дихання насіння, яке проростає, дужа висока
(Овчаров, 1962).

Виключно висока біологічна активність жиророзчинних вітамінів
і їх ліпофільна природа разом забезпечують можливість проникнення і
вбудову в ліпідну фазу біомембрані тому ці вітаміни можуть грати важливу
роль в їх функціонуванні (Спиричев,Конь,1978).

І. Участь вітаміну С в різних фізіологічних процесах в рослинних
організмах.

Вітамін С бере участь у фотосинтезі. Вітамін С бере участь у
фотовідновленні хлорофілу. Цей процес можна зобразити на схемі.

Схема 1: участь аскорбінової кислоти у відновленні хлорофілу.

АН2+ +О2

АН + ХН Х ХО2 Х + ВО2

+ О2 О2

А + Н2О Х + Н2О Х + Н2О2 + А

Н2О2 О2

де АН – аскорбінова кислота, А – дегідроаскорбінова кислота, В –
олеїнова кислота, “Х” – сенсибілізований хлорофіл.

Було помічено, що більша кількість аскорбінової кислоти
накопичується в листках південної крони. Ці факти вказують на те, що під
час фотосинтезу синтезується аскорбінова кислота. Отже вона не тільки
впливає на фотосинтетичні процеси, а й сама виникає під час їх
проходження. Крім цього відомі факти, що аскорбінова кислота включається
в процеси, що проходять після фотолізу води, при фотосинтетичному
фосфорилюванні.

Також аскорбінова кислота сильно пов’язана з процесами дихання
рослин. Аскорбінова кислота має властивості оборотньо-окисно-відновної
системи, яка здатна бути акцептором і донором водню.

При введенні аскорбінової кислоти в тканини рослин, посилюються
інтенсивність дихання цих рослин. Аскорбінова кислота в рослинах і її
участь в диханні надає велику стійкість організмам, так як вона
окислюється завдяки різним “кінцевим” оксидазам, тобто функціонує в
різних умовах температури й на різних етапах розвитку рослин.

Аскорбінова кислота бере участь в мінеральному живленні рослин.

Як уже відомо при добавлянні окремих мінеральних добрив на бідних
грунтах у рослинах відмічається збільшення вмісту аскорбінової кислоти.
На грунтах з великим вмістом різних елементів мінерального живлення
добавляння добрив ефекту не дає (Спірідонова, 1968). Є такі елементи,
при внесенні яких у грунт, вміст аскорбінової кислоти в органах рослин
понижується. Таким елементом є азот. Часто при добавленні азоту
спостерігається пониження концентрації аскорбінової кислоти.

Особливо негативно діє добавлення азоту на вміст аскорбінової
кислоти в органах плодових дерев (Спірідонова, 1968). Але внесення бору
і мангану збільшує вміст аскорбінової кислоти в листках і коренях
петрушки, пастернаку та інших рослин. Цинк і молібден також впливають на
збільшення вмісту вітаміну С в різних органах рослин.

Аскорбінова кислота також бере участь у такому важливому
біологічному процесі як водообмін. Це доказано, наприклад,
експериментом, який провели Н. С. Спірідонова і її співробітники над
кукурудзою і картоплею у 1968 році. Вони показали, що під дією
аскорбінової кислоти посилюється транспірація у рослин, а також
швидкість руху води по рослині, так, як аскорбінова кислота сприяє
прискоренню цілого ряду метаболічних процесів.

Крім цих усіх аскорбінова кислота бере участь у пристосувальних
процесах до умов навколишнього середовища (Спірідонова, 1968).

Це підтверджено дослідженнями, що проводились на різних висотах
над рівнем моря. Виявили, що в тих рослинах, що ростуть на більших
висотах накопичується більше аскорбінової кислоти. Це можна пояснити
посиленням фотосинтетичних процесів у рослин і окисної активності
тканин, у рослин при підйомі вгору.

II. Вітамін В1. Його роль для фізіологічних і біохімічних процесів
рослин.

Цей вітамін входить до коферменту деяких ферментів, наприклад
піруватдекарбоксилази, яка відщеплює СО2 від піровиноградної кислоти,
цим самим грає важливу роль в перетворенні вуглеводів в організмі
рослин, бактерій і тварин (Пивоваренко,1995).В склад ферментів вітамін
В1 входить у вигляді фосфорнокислого ефіру тіамінпірофосфату.
Фізіологічне значення тіаміну теж велике. Наприклад, тіамін має
стимулюючу дію на ріст корінців багатьох рослин, на дихання рослин
(Овчаров, 1962), у процесі фотосинтезу та інших фізіологічних процесах
(Кретович, 1986).

Значення вітаміну В1 для росту коренів показують експерименти з
ізольованими корінцями. Ріст ізольованих верхівок коренів вищих рослин
спостерігається тільки при внесенні в живильне середовище вітаміну В1.
Корені деяких рослин (наприклад, помідорів) потребують внесення в
живильне середовище не тільки вітаміну В1, але й пірідоксину. В ряду
випадків ріст коренів в штучних умовах спостерігається лише при внесенні
разом з вітаміном В1 і пірідоксином ще й нікотинової кислоти (Рубін,
1976).

Ці спостереження показують, що корені вищих рослин можуть
синтезувати більшість вітамінів, але не можуть синтезувати вітаміну В1,
який необхідний для їх росту, а деякі інші не синтезують нікотинову
кислоту і пірідоксин. Але ці вітаміни синтезуються в інших місцях і
поступають в корені через провідні тканини. Також вітамін В1 важливий
для зелених листків (Кудряшов, 1953). Гершенович і Микитіна показали,
що тіамін володіє здатністю гальмувати окислення аскорбінової кислоти,
тобто являться сильним стабілізатором її окислення в клітинному
метаболізмі.

ІІІ. Фізіологічна роль вітаміну В2 в життєдіяльності рослин.

Вітамін В2 синтезується тільки в рослинах і вдеяких
мікрорганізмах. На світлі і в темряві утворюється з однаковою швидкістю
(Плешков,1969).

Рибофлавін в сполуці з фосфорною кислотою входить в склад ряду
ферментів, які відіграють важливу роль в обміні речовин. В складі
флавінмононуклеотиду (ФМН) входить в склад окисно-відновних ферментів.

Ці ферменти беруть участь в переносі атомів Гідрогену. В
рослинних організмах як і в організмах тварин рибофлавін міститься як у
вільному вигляді, так і у вигляді фосфатного ефіру. Вітамін бере участь
у диханні рослин, входячи в склад ферментів дихання, які беруть участь в
окислювальному декарбоксилуванні піровиноградної кислоти. Вітамін В2
входить до коферменту ФАД, який входить до ліпоаміддегідрогенази.
(Кретович, 1986., Пивоваренко,1995). Також необхідний цей вітамін для
проростання пилку разом з вітаміном В1 і С (Кудряшов, 1953). Вміст цього
вітаміну залежить від мінерального живлення і особливо падає при
недостачі азоту і магнію в грунті або в поживному середовищі
(Плешков,1965).

IV. Фізіологічна і біологічна роль вітаміну В6 для рослин.

Альдегідне похідне вітаміну В6 в вигляді фосфатного ефіру входить
в склад ферментів, які каталізують різні перетворення амінокислот: їх
декарбоксилювання, реакції переамінування (Березовський,1959).

При авітамінозі В6 відмічають глибоке порушення в синтезі і обміні
триптофану. Вітамін В6 може в деяких випадках посилювати дію вітаміну В1
Бере участь у процесі фотосинтезу та інших біологічних і фізіологічних
процесах (Овчаров, 1962).

V. Фізіологічна роль вітаміну РР.

Ця речовина у рослинах знаходиться, головним чином, у вигляді
кислоти, яка перетворюється в нікотинамід і у його вигляді входить в
склад окисно-відновних ферментів – дегідрогеназ, які каталізують
відбирання атомів Гідрогену від органічних речовин, які при цьому
окислюються. Далі ці атоми Гідрогену дані ферменти передають
окисно-відновним ферментам, в склад яких входить рибофлавін. Також
входячи до цих ферментів вітамін РР бере участь у бродінні і диханні
рослин (Кретович, 1986). Також нікотинова кислота є стимулятором росту
рослинного організму (Труфанов, 1959). Інтенсивний синтез нікотинової
кислоти в рослинах починається одночасно з пророрстанням насіння.
Наприклад, при проростанні насіння квасолі вміст вітаміну через три дні
підвищується від 1,9 до 8,9мг. Синтез нікотинової кислоти прискорюється
на світлі (Плешков,1965).

VI. Фізіологічна роль пантотенової кислоти.

Входить до складу коферменту А (коензиму А), при участі якого
проходить активування й перенесення залишків оцтової кислоти і інших
кислотних залишків (ацилів), синтез лимонної кислоти, жирних кислот,
стеролів, гліцеридів і інших сполук.

На ізольованих зародках гороху також було показано, що і
пантотенова кислота безумовно потрібна для рослини, що розвивається.
Проростки люцерни, що вирощуються на стерильному поживному середовищі,
подібно гороху, розвиваються значно краще при добавлянні в поживне
середовище незначної кількості пантотенової кислоти (Кудряшов, 1953)

VIІ. Значення вітаміну Н (біотину) для рослинних організмів.

Вітамін Н бере участь в перетвореннях деяких амінокислот
(аспарагінової кислоти, серину, треоніну). Біотин входить в склад
активної групи ферментів які каталізують процеси карбоксилювання жирних
кислот, тобто приєднання СО2, яке супроводжується подовженням
вуглеводневого ланцюга жирних кислот. Також біотин грає роль у
приєднанні СО2 до ацетилкоферменту А і утворенні малонілкоферменту А.

О О

СН3 – С + СО2 НООС – СН2 – С

S – C0A S – C0A

Ацетил кофермент А малоніл кофермент А. (Кретович, 1986

По деяким даним один біотин стимулює переважно ріст стебла і майже не
впливає на ріст кореня. По іншим даним біотин потрібен для росту кореня.
Наприклад, якщо для культури ізольованих тканин в поживне середовище
добавляти вітамін В1, а біотин не вносити, то ріст стебла
уповільнуюється. Коли ж вносити оба вітаміни, тоді накопичення сухої
речовини в порівнянні з піддослідними рослинами збільшується на 60 %
(Кудряшов, 1953).

VIІІ. Роль фолієвої кислоти (В9) в життєдіяльності рослин

Вітаміни групи фолієвої кислоти беруть участь в цілому комплексі
реакцій синтезу і переносу одновуглецевих сполук, фіксації СО2
синьо-зеленими водоростями, в реакціях метилювання, перенесення
активного
формальдегіду
(Андреєва, 1963).

В вільному вигляді фолієва кислота в живих організмах може служити
активатором протонів при цьому бере участь в окисно-відновних реакціях.
Відновлення фолієвої кислоти проходить за участю НАДФН-Н+ , а фермент,
що каталізує відновлення фолату, називається
7,8-дигідрофолат-НАДФ-оксидоредуктазою. Подальше відновлення
дигідрофолата в тетрагідрофолат здійснюється теж за участю НАДФН-Н+-
коферменту. Фермент називається
5,6,7,8-тетрагідрофолат-НАДФ-оксидоредуктаза. Основна біохімічна роль
фолієвої кислоти виконується тетрагідрофолієвою кислотою, яка здатна при
сполученні з відповідними апоферментами до переносу одно вуглецевих
радикалів:

Н–С=О; CН2ОН; –СН3; – СН2 – ;
НС =NH.

| |
|

форміл оксиметил метил
метилен форміміно.

Процес перенесення формільних залишків, який проходить за участі
води, є основною функцією тетрагідрофолієвої кислоти в організмі і
називається трансфoрмілуванням (Плешков,1969). Внаслідок перенесень
таких залишків фолієва кислота бере участь у взаємоперетворенні
амінокислот серина і гліцина.

Також фолієва кислота бере участь у синтезі пуринових основ, а,
як наслідок, нуклеїнових кислот (Колотилова, Глушанков, 1976).

IХ. Фізіологічна роль вітамінів групи Q (убіхінони) для рослин.

Вони зосереджені у внутрішніх мембранах мітохондрій, ядрах
апараті Гольджі. Мають ліпідоподібну структуру.

У рослинах вони беруть участь у фотосинтезі і стабілізують
клітинні мембрани, захищаючи ненасичені залишки жирних кислот ліпідів
від окислення.

Отже потрібно сказати, що вітаміни грають дуже важливу роль в
біохімічних та фізіологічних процесах рослин, тому рослини повинні їх
синтезувати. Ми бачили, що при добавлянні пантотенової кислоти в поживне
середовище де вирощували ізольовані зародки гороху і люцерни, ці зародки
почали рости скоріше. Це говорить, що для розвитку зародків і для
проростання насіння взагалі потрібні вітаміни, бо вони грають важливу
роль, як в житті сформованої рослини, так і в процесах, які проходять
під час проростання насіння рослин.

Х.Фізіологічна роль вітамінів групи К для рослин.

В рослинах вітаміни групи К беруть участь в окисно-відновних
процесах і, в тому числі, в процесі фотосинтетичного фосфорилювання, де
вони служать проміжними переносниками електронів. Тому і біосинтез
вітаміну К зв`язаний з біосинтезом хлорофілу, а також тому в
хлоропластах цього вітаміну більше, ніж у цитоплазмі клітини
(Плешков,1965).

ХІ. Роль вітаміну Е для життєдіяльності рослин.

Токофероли запобігають окисленню і прогірканню рослинних масел,
тобто являються антиокисниками (Кретович, 1986). Бере участь у багатьох
видах обміну у тваринних організмів (Дунаєвський, 1990) і, можливо,
рослинних організмах. Цей вітамін представлений чотирьма ізомерами, які
мають назву Ь-, в-, і г- токоферолів. Вітамін Е, подібно іншим вітамінам
, здатен вступати в сполуку з білком, причому етерифікований вітамін
більш сильно зв‘язаний з ним, ніж вільний, так як відокремлюється лише
при денатурації (Захарова,1954).

Найбільше міститься в зародках злаків і зелених листках, значить
тут він грає найважливішу для рослин роль. Як правило, у зелених листях
його більше ніж у стеблі і інших органах, а у всіх зелених частинах
рослини його до 67% порівняно з зеленими частинами
(Луцевська,Савінов,1953). Вітамін Е стимулює цвітіння дводольних
рослин, бо по хімічній будові належить до класу ізопреноїдів які
необхідні для цвітіння (Рубін, 1967).

ХІІ. Значення провітамінів А (каротиноїдів) для життєдіяльності
рослинних організмів.

У рослинах є лише попередники вітаміну А – каротини. Вони
поширені в листках і особливо в плодах і грають важливу роль в
обміні речовин в епітеліальних і інших клітин рослини
(Березовський,1959). Каротини є пігментами рослин, вони приймають участь
у фотохімічних реакціях при фотосинтезі (Кретович, 1986). Також каротин
приймає участь в регуляції окислювальних процесів в рослинній клітині.
Це підтверджує наявність оксидази каротину в рослинах і гальмування
каротинами в рослинних клітинах каталітичного окислення аскорбінової
кислоти аскорбінооксидазою. Очевидно, між цими двома системами
аскорбінова кислота – аскорбінооксидаза і каротин – каротиноксидаза є
рівновага, яка має значення в окислювальних і дихальних процесах
рослинної клітини (Труфанов, 1959).

ЛІТЕРАТУРА

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эффэкт витаминов группы B при действии гербицида 2,4-Д на ультра
структурную организацию клеток растительного организма //Физиология и
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2. Гаврыш С.Ф. Томаты. – М.: Госсельхозиздат, 1887. – 71с.

3. Гершенович З.С., Микитина А.И. Взаимодейсвие тиамина и аскорбиновой
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4. Дунаевский Г.А., Попик С.Я., Овощи и фрукты в питании здорового и
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10. Овчаров К.Е. Физиологические основы всхожести семян. – М.: Наука,
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13. Плешков Б.П. Биохимия сельскохозяйственных растений /под ред.акад.
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22. Строганов Б.П., Генкель К.П. Роль витаминов в жизнедеятельности
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23. Труфанов А.В. Биохимия и физиология витаминов и антивитаминов. – М.:
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Також використовували Біологічний енциклопедичний словник. (Електронну
версію).

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