10강. 종자의 생물학과 중요성

1. 종자의 해부구조

2. 종자의 화학적 조성

3. 종자의 발아

4. 종자의 유전학

1. 종자의 구조

Ovary (씨방; 자방) Seed (종자)Ovule (배주)난세포(egg cell) + 웅핵(sperm nuclei) ---> 2 ⓝ ---배(embryo)

ⓝ ⓝ극핵(polar nuclei) + " ---> 3 ⓝ ---------배유(endosperm)

2 ⓝ 3배체

Integument(주피) 2 ⓝ 종피(seed coat)

Ovary wall(자방벽) 2 ⓝ 과실(fruit)

과피 (pericarp; fruit coat )

배 (Embryo)

(상배축)

(외배엽)

(3본엽)

(배반)

(생장점)

(자엽초)

(중배축)

(근초)

(배축)

(근관)

(배축)

(자엽)

(전형성층)

냉이

배유 (Endosperm)

벼 종자 횡단면

벼 종자의 배유층

콩 종자 횡단면

종피 (Seed coat)

벼

(주심조직)

(주피조직)

2-3DAP

5-6DAP

15DAP

matured

목화

Ovule

콩종피

(하피)

(망상맥)

벼 aleurone layer(호분층)

등부분Multi-layered

aleurone layer

종자의모양

장미과 뽕나무과 메꽃과 미모사과

백합과

범의귀과 난초과

사초과 포아풀과(화본과)

미나리아재비과

수련과

명아주과

2. 종자의 화학적 조성

전분 단백질 지방질

벼 64.3 7.9 1.8

밀 69.7 13.2 1.9

옥수수 63.6 8.2 3.9

보리 55.8 8.7 1.9

완두 50.0 23.4 1.2

녹두 35.6 23.6 0.2

콩 23.5 37.9 18.0

땅콩 5.9 30.4 47.7

유채 - 20.4 43.6

해바라기 - 16.8 25.9

(1) 탄수화물

a. 전분(starch) : amylose, amylopectinb. Hemicellulose: xylan, mannan, galactan 등c. Others

- 점액질—종자 표면: pectin화합물, polyuronide, galacturonide 등

- Pectin 화합물-- 세포벽과 중륵의 구성성분: pectic acid, pectin,protopectin 등

(2) 저장단백질(단순단백질) a. Albumin : water-solubleb. Globulin : water-insoluble, dilute salt-solublec. Glutelin : water/alcohol/salt-insoluble, weak acid/alkali-solubled. Prolamin : water/salt-insoluble, weak acid/alkali-soluble,

pepsin-resistant(복합단백질)

prolamin

globulin/glutelin

globulin

prolamin

* Triticin: a carbohydrate resembling irisin

*

Osborne fraction

Solubility behaviour Composition Biological role Functional role밀 단백질 중 비율(%)

AlbuminWater and dilute buffers

Non-gluten proteins (mainly monomeric)

Metabolic and structural proteins

Protection from pathogens

15~20

Globulin Dilute saltProviding food reserve to embryo

Gliadin Aqueous alcohols

Gluten proteins (mainly monomeric gliadins and low molecular weight glutenin polymers) Prolamins-type

seed storage proteins

Dough viscosity/plasticity

40~45

Glutenin Dilute acetic acid Gluten proteins (mainly HMW gluteninpolymers)

40

Residue

Unextractable in water and dilute buffers but extract-able with Urea+ DTT+SDS SDS+ Phosphate buffers+ sonication etc

Gluten proteins (high molecular weight polymers) and polymeric non-gluten proteins (triticins)

Prolamins-type (gluten) and globulin-type (triticin) seed storage proteins

Ali Hafeez Malik, 2009, ISSN 1654-3580

Classification of proteins according to Osborne, modified from Goesaert et al., (2005)

밀의 단백질 함량: 10~15%

* 단백질 Gluten

강력분: 글루텐 13% 이상 제빵중력분:10~13% 제면, 제빵박력분: 10% 이하 제과, 튀김가루

Gluten ( glue + protein) : 밀, 보리, 호밀에도 많이 존재

1. 점성의 단백질2. 소화 잘 안됨3. Gliadin이 소장의 점막으로 흡수되어 문제를 일으킴 – 인구의 약 10%

밀가루에 물을 넣고 반죽하면 이 두단백질이 섞이면서 단백질 네트워크를형성하는데, 이를 글루텐.

탄성점성신장성

심한 경우celiac병

(3) 지방질- 지방산(fatty acid)- glycerol과 기타 알콜류

: 자방산과 결합하여 다양한 지방질

Lauric Myristic Palmitic Stearic Oleic Linoleic

ω-6

Linolenic

ω -3

코코넡

올리브

땅콩

옥수수

콩

해바라기

현미

참깨

45.0

-

-

-

-

-

-

-

20.0

-

-

-

-

-

-

-

5.0

14.6

8.5

6.0

11.0

3.5

15.0

9.5

3.0

-

6.0

2.0

2.0

2.9

1.9

4.5

6.0

75.4

51.6

44.0

20.0

33.4

42.5

42.5

-

10.0

26.0

48.0

64.0

57.5

39.1

42.5

-

-

-

-

3.0

-

-

1.0

종자의 지방산함량(%)

포화지방산

흡수후 불포화과정 거쳐 변함

(4) 그외성분들

a. Tannin: 종자의과피에주로분포

b. Alkaloids: 담배의 nicotine, 양귀비의 morphine 등

c. Glucoside: sinigrin (흑겨자) 등

d. Phytates: phytin 등

e. Hormones: auxin, gibberellin, cytokinin 등

f. Vitamins

쌀의 부위별주요 영양소와 미네랄

Rough rice = hull + brown rice (bran + (polish) + white rice)

(pericarp + seed coat + nucellus + aleurone layer + ~endo.)

쌀의부위별비타민함량 (ug/g)

Bran contains more of the pericarp, seed coat, nucellus, aleurone layer and germs than polish,

Polish: more starchy endosperm than bran

쌀의부위별미네랄함량

Nutrients: Content per 100 g

Energy 1364 kJ (322 kcal)

Water 8.5 g

Protein 33.7 g

Lipids 18.1 g

Carbohydrate 6.1 g

Minerals 4.7 g

Minerals

Sodium 4 mg

Potassium 1740 mg

Magnesium 245 mg

Calcium 255 mg

Manganese 3 mg

Iron 8.6 mg

Copper 110 µg

Zinc 1 mg

Phosphorus 590 mg

Chloride 7 mg

Iodine 6 µg

Selenium 60 µg

Vitamins

Carotene 380 µg

Vitamin E 1500 µg

Vitamin K 190 µg

Vitamin B1 990 µg

Vitamin B2 520 µg

Nicotinamide 2510 µg

Pantothenic acid 1920 µg

Vitamin B6 1190 µg

Biotin 60 µg

Folic acid 230 µg

Amino Acids

Arg 2360 mg

His 830 mg

Ile 1780 mg

Leu 2840 mg

Lys 1900 mg

Met 580 mg

Phe 1970 mg

Thr 1490 mg

Trp 450 mg

Tyr 1250 mg

Val 1760 mg

Carbohydrates

Glucose 5 mg

Sucrose 6100 mg

Lipids

Palmitic acid 1580 mg

Stearic acid 590 mg

Oleic acid 3790 mg

Linolic acid 8650 mg

Linoleic acid 1000 mg

Other

Lecithin 1.1-3.2 g

Purines 380 mg

Reference: Deutsche Forschungsanstalt für Lebensmittelchemie, Garching bei München (ed), Der kleine "Souci-Fachmann-Kraut" Lebensmitteltabelle für die Praxis, WVG, Stuttgart 1991

Nutritional composition of soybean seeds

종자의 휴면

major mechanisms for imposing dormancy:

1. Seed coverings : Restrict water uptake, embryo expansion,

gas permeability, leaching of inhibitors

2. Chemical inhibitors include growth regulators etc.

3. Morphological aspects such as small, undeveloped

embryos.

3. 종자의 발아

(Stratification)

종자의 발아

상처

Effect of fungal species on germination of Opuntia leucotricha seeds

Photomicrographs of fungal hyphae eroding the endocarp of seeds, obtained with an environmental scanning electron microscope:(A) Penicillium chrysogenum, (B) Phoma sp., (C) Trichoderma koningii and (D) control (sterile seed). Plant Biology 13 (2011) 154–159

Alternaria and Aspergillus fungi increase germination of Astragalus utahensis

Fungi 와 같이 후숙 처리시 발아율 제고 효과를 본 예)

(GA대신에)

(광발아종자)

(in each cell)(적색광)

(근적색광)

(종피)

(배축)

(유아)

(제; 배꼽)

(떡잎;자엽)(주공)

(하배축)

(상배축)

(근초)

(유근)

(엽초)

(초엽)

(과피)

(배유)

(유아)

(부정근)

4. 종자의 유전학

(1)형태/크기

Gene Trait Protein function properties

Ref Years

*GS3 GL,TGW

Transmembrane protein, putative G-protein gamma subunit

NR Fan 2006

*qSW5 GW Putative nuclear protein related ubiquitinproteasome pathway

NR Shomura

2008

*GW2 GW RING-type E3 ubiquitin ligase NR Song 2007

*GS5 GW,GF

Putative serine carboxypeptidase belong topeptidase S10 family

PR Li 2011

*GW8/OsSPL16

GW SQUAMOSA promoter-binding protein-like 16 PR wang 2012

TGW6 TGW,GL

Iindole-3-acetic acid (IAA)-glucose hydrolase NR Ishimaru

2013

GS6 GW,TGW

a member of the GRAS gene family NR Sun 2013

벼 종자 크기 및 모양에 관여하는 것으로 보고된 유전자들

: 주동유전자(major genes), 미동유전자(minor genes)

Os05g0187500) – chr. 5

(g) Association analysis of more than 100rice cultivars. Cultivars were categorized into three groups by allele polymorphisms in the qSW5 FNP. (Shomura et al., 2008)

5367k

Os05g0187500

4.4kb

1212 deletion

950 deletionNipponbare typeIndica Ⅱ typekasalath type

Deletion size

197

A

16729k 16736k

C

→

Nipponbare

Sona

A-allele C-allele

45bp

Os03g0407400

(Fan등, 2006)

유전자의 효과 예)

(2) 이화학적 성분 함량 :

Purple pericarp (anthocyanin) PpPpPbPb Purple

ppppPbPb BrownPpPppbpb , pppppbpb White

Red pericarp (tannin) RcRcRdRd Purple-Red

RcRcrdrd BrownrcrcRdRd , rcrcrdrd White

Wxa

Wxb

Wxop

이화학성분의대부분은양적형질유전자에의해 결정된다.

: 주동유전자(major genes), 미동유전자(minor genes)

Anthocyanin pathway in maize

(BMC Plant Biology 2013, 13:42)

C1, P : transcrition factorsPAL1: Phenylalanine Ammonia Lyase 1C2: chalcone synthaseChi: chalcone isomeraseF3H: flavon-3-hydroxylaseA1, A2: anthocyanin1, 2Bz1, 2: bronze 1, 2

Jirin/LK1A-2-12-1-1

05

10152025303540

0.00

170.00

340.00

510.00

680.00

850.00

1020

.00

1190

.00

1360

.00

1530

.00

1700

.00

1870

.00

2040

.00

2210

.00

c3g content

Num

ber of

lines

Heugjunju/Hwacheong ges

01020304050607080

0.00

50.00

100.00

150.00

200.00

250.00

300.00

350.00

400.00

450.00

500.00

550.00

600.00

c3g content

Num

ber of

lines

No2/Heugbal

01020304050607080

0.00

50.00

100.00

150.00

200.00

250.00

300.00

350.00

400.00

450.00

500.00

550.00

600.00

c3g content

Num

ber of

lines

Jirin/Heidao38

05

10152025303540

0.00

200.00

400.00

600.00

800.00

1000

.00

1200

.00

1400

.00

c3g content

Num

ber of

lines

Fig 2. Segregation of C3G concentration (mg/100g brown rice) of rice grain in F2 of four crosses

벼유색미에서Anthocyanin 함량의유전분리양적형질

(Hexi 41 (P1)/

Ilpumbyeo (P2))

0

10

20

30

-10 -7 -4 -1 2Greenness (a)

No.

of

indiv

idua

l

P1 P2

P2

(Hexi 41 (P1)/

Hwachung wx sgr (P2))

0

10

20

30

-10 -7 -4 -1 2Greenness (a)

No.

of

indiv

idual

P1 P2

(Hex 41 (P1)/

Dasanbyeo (P2))

0

15

30

45

60

-10 -7 -4 -1 2Greenness (a)

No.

of

indiv

idua

l

P1 P2

(Jilinluimi-1 (P1)/

Ilpumbyeo (P2))

0

10

20

30

40

-10 -7 -4 -1 2Greenness (a)

No.

of

indiv

idua

l

P1

P2

(Jilinluimi-1 (P1)/

Hwachung wx sgr (P2))

0

10

20

30

40

-10 -7 -4 -1 2Greenness (a)

No.

of

indiv

idua

l

P1

P2

(Beijingluimi-1 (P1)/

Ilpumbyeo (P2))

0

10

20

30

40

-10 -7 -4 -1 2 Greenness (a)

No.

of

indiv

idua

l

P1

P2

그림. 녹색쌀과 일반미간, 녹색쌀과 녹색쌀간의 F2 조합에서의 녹색도의 분리양상

* 곡류에서 배유와 배의 유전자형

메벼품종

찰벼품종

WxWx

WxWxWx

(3n)

wxwx

wxwxwx

(3n)x

Wxwx

WxWxwx

(3n) 메벼F1

wxwxwx WxWxWx

WxWxwx

X =

wxwxwxWxWxWx

Wxwxwx

X =

(배유의 유전자형)

wxwxwx X wxwxwx wxwxwx=

WxWxWxWxWxWx =WxWxWxX

(아밀로스함량, 예)

0 %

10 %

16 %

19 %

Wx gene 의 dosage 효과