Fern spore germination in response to environmental factors

doi:10.1007/s11515-015-1342-6

Front. Biol.

2015, Vol. 10

Issue (4) : 358-376 https://doi.org/10.1007/s11515-015-1342-6

REVIEW

Fern spore germination in response to environmental factors

Jinwei Suo^1,²,Sixue Chen³,Qi Zhao¹,Lei SHI^4,^*(

),Shaojun Dai^1,^*(

)

1. Development Center of Plant Germplasm Resources, College of Life and Environmental Sciences, Shanghai Normal University, Shanghai 200234, China
2. Alkali Soil Natural Environmental Science Center, Northeast Forestry University, Key Laboratory of Saline-alkali Vegetation Ecology Restoration in Oil Field, Ministry of Education, Harbin 150040, China
3. Department of Biology, Genetics Institute, Plant Molecular and Cellular Biology Program, University of Florida, Gainesville, FL 32610, USA
4. Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China

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Abstract

Fern spore germination gives rise to the rhizoid and protonemal cell through asymmetric cell division, and then develops into a gametophyte. Spore germination is also a representative single-cell model for the investigation of nuclear polar movement, asymmetrical cell division, polarity establishment and rhizoid tip-growth. These processes are affected by various environmental factors, such as light, gravity, phytohormones, metal ions, and temperature. Here, we present a catalog of spore germination in response to different environmental factors. They are as follows: (1) Representative modes of light affecting spore germination from different fern species include red light-stimulated and far red light-inhibited spore germination, far red light-uninhibited spore germination, blue light-inhibited spore germination, and spore germination in the dark. The optimal light intensity and illumination time for spore germination are different among various fern species. Light response upon spore germination is initiated from the cell mitosis that regulated by phytochromes (PHYs) and cryptochromes (CRYs). AcPHY2, AcCRY3 and/or AcCRY4 are hypothesized to be involved in spore germination; (2) Gravity and calcium are crucial to early nuclear movement and polarity establishment of spores; (3) Gibberellin and antheridiogen can initiate and promote spore germination in many species, but abscisic acid, jasmonic acid, and ethylene pose only minor effects; (4) Spores can obtain the maximal germination rate in their favorable growth medium. Moreover, metal ions, pH, and spore density in the culture medium also affect spore germination; (5) Most fern spores germinate at 25°C, and an optimal CO₂ concentration is necessary for spore germination of certain fern plants. These provide valuable information for understanding fern spore germination in response to environmental factors.

Keywords fern spore germination environmental factors

Corresponding Author(s): Lei SHI,Shaojun Dai

Just Accepted Date: 26 December 2014 Online First Date: 28 January 2015 Issue Date: 14 August 2015

Cite this article:

Qi Zhao,Lei SHI,Shaojun Dai, et al. Fern spore germination in response to environmental factors[J]. Front. Biol., 2015, 10(4): 358-376.

URL:

https://academic.hep.com.cn/fib/EN/10.1007/s11515-015-1342-6
https://academic.hep.com.cn/fib/EN/Y2015/V10/I4/358

Fig.1 Phylogeny depicting relationships of fern species whose spore germination is affected by various environmental factors. The effects of environmental factor on spore germination were shown in red (promotion), green (inhibition) and blue (no influence). ABA: Abscisic acid; An: Antheridiogen; B: Blue light; B-R: Blue light followed by red light; CO_2a: CO₂ treatment concentration for the spores of P. piloselloides (219–3360 cm³·m^-3) and O. sensibilis (0–2%); CO_2b: CO₂ treatment concentration for O. sensibilis spores (5%–15%); D: Darkness; E: Ethylene (≥1μL·L^-1); E-im: The treatment of ethylene on the immature fern spores; E-m: The treatment of ethylene on the mature spores; FR: Far red light; FR_＞24h-R: 24h far red light treatment followed by red light; FR(B)-B(FR): Far red light followed by blue light or blue light followed by far red light; FR-R: Far red light followed by red light; G: Green light; GA: Gibberellin acid; GA₄₊₇: A kind of gibberellin acid; Gr: Gravity; I₁: Ca²⁺, Mg²⁺, or Mn²⁺; I₂: La³⁺ or Cd²⁺; ID₁: Inoculation density, 50 spores·mL^-1; ID₂: Inoculation density, 3000 spores/cm²; JA: Jasmonic acid; LD₁: Long time of light irradiation (≥24 h); LD₂: Short time irradiation (<24 h); LI-h: High light intensity; LI-l: Low light intensity; LI-s: Leaf shade; LI-w: White light; MS₁: Medium with lower concentration (<1/2MS: 1/4MS, 1/5MS, 1/10MS etc.); MS₂:1/2 MS; MS₃: MS or≤1/2MS; MS₄: The liquid and solid medium; N₁: Other nutrition in the medium: such as, sulphite, oxybarbiturates, lipophilic solvents, and the leachates of Eupatorium adenophorum; N₂: Other nutrition in the medium: such as, thiamine, pyridoxine, glycine, glutamate, kinetin, and extract of P. aquilinum leave; pH₁: A wide pH range (3.7 to 9.7 or 4.5 to 8.5); pH₂: 3.7 to 6.7; pH₃: 7 to 8; R: Red light; R-B: Red light followed by blue light; R/FR-B_6-72h: Red or far red light followed by blue light for 6–72 h; R-D_3-5h-B/FR: Red light followed by blue or far red light after an intervening dark period (3–5 h); R-FR: Red light followed by far red light; R-FR-R_s: Red light followed by far red light, and then red light for short time (5 min); R-FR-R_l: Red light followed by far red light, and then red light for 8 h; R_38h-B/FR-X : 38 h red light treatment followed by blue or far red light, and then a series of brief red, far red, and blue light irradiation; (R-)B-R_h/l: (Red light-) Blue light followed by red light of high intensity (1–2 min) or long time (1h); (R-)B-R_h-B_s: (Red light-) Blue light followed by red light with high intensity (2 min), and then blue light for short time (1–2 min); (R-)B-R_h- B_l/FR: (Red light-)Blue light followed by red light with high intensity (2 min), and then blue light for long time (1–2 h) or far red light; S-L: Low concentration of sucrose in MS; S-H: High concentration of sucrose in MS; T<15: The optimum germination temperature less than 15 °C; Top-20: The maximum germination rate occurred at 20 °C; Top-25: The optimum germination temperature is 25 °C; Top-30: The maximum germination rate occurred at 30 °C. W: White light; Y: Yellow light.

Environmental factors	Plant species	Treatment condition(concentration, time)	Effects on spore germination	References
Red light	Adiantum capillus-veneris	5 min (followed in darkness for 168 h)	P (58.3%)	Furuya et al., 1997
	38 h (followed in darkness for 38 h)	P (78.3%±2.6%)
	Alsophila metteniana	12 h daily	P (65.66%±4.04%)	Du et al., 2009a
	Asplenium nidus	≥72 h	P (initiated both rhizoid and protonema)	Raghavan, 1971
	12 h	Only formed rhizoids
	Cheilanthes farinosa	30 s	P (80%)	Raghavan, 1973
	Cheilanthes feei	–	P (88%)	Nondorf et al., 2003
	Lycopodium clavatum	30 min daily	I (5.9%)	Whittier, 2008
	Lygodium japonicum	–	P	Sugai et al., 1977
	Mohria caffrorum	144 h or 12 h or 24 h	86.2%±2.7% or 42.3%±3.3% or 79.1%±3.9%	Reynolds and Raghavan., 1982
		5 min/h (during a 24 h period)	SI (77.6%±1.3%)
		5 min/4 h (during a 24 h period)	I (39.4%±2.8%)
		5 min/8 h (during a 24 h period)	I (15.4%±0.7%)
	Onoclea sensibilis	5 min	P (63.2%±2.5%)	Towill et al., 1973
	Ophioglossum crotalophoroides	20 min daily	I (0%)	Whittier, 2006
	Polypodium aureum	1 d, 3 d, 5 d	33%, 61%, 79%	Spiess et al., 1977
	Pteris vittata	–	P	Sugai et al., 1967
	Thelypteris kunthii	8 h or 18 h	P (34.8% and 46.5%±2.05%)	Huckaby and Raghavan, 1981
		5 min/1 h for 18 h	I (37.2%±2.63%)
		5 min/3 h for 18 h	I (32.9%±2.17%)
		R 8 h, D 24 h	P (34.6%±4.45%)
Far-red light	Adiantum capillus-veneris	5 min (followed in darkness for 168 h)	I (1.2%)	Furuya et al., 1997
	Cheilanthes feei	–	P	Nondorf et al., 2003
	Lycopodium clavatum	30 min daily	P (95.8%)	Whittier, 2008
	Lygodium heterodoxum	12 h daily	P (74%)	Pérez-García et al., 1994
	Mohria caffrorum	144 h	P (84.0%±3.1%)	Reynolds and Raghavan, 1982
		12 h	P
		24 h	P (80.6%±1.2%)
		5 min/h (during a 24 h period)	P (78.8%±1.9%)
		5 min/4 h (during a 24 h period)	I (35.2%±3.3%)
		5 min/8 h (during a 24 h period)	I (9.0%±2.6%)
	Onoclea sensibilis	5 min	P (75.2%±3.1%)	Towill et al., 1973
	Ophioglossum crotalophoroides	20 min daily	P (25.3%)	Whittier, 2006
Darkness	Adiantum capillus-veneris	168 h or 38 h	I (0%)	Furuya et al., 1997
	Alsophila metteniana	–	I (0%)	Du et al., 2009a
	Thelypteris kunthii	18 h or 24 h	I (0%)	Huckaby and Raghavan, 1981
	Cheilanthes farinosa	4 weeks	I (8%)	Raghavan, 1973
	Cheilanthes feei	–	I (7.7%)	Nondorf et al., 2003
	Lycopodium clavatum	24 h daily	P (98.7%)	Whittier, 2008
	Mohria caffrorum	144 h	I (0%)	Reynolds and Raghavan, 1982
	Onoclea sensibilis	6 d	I (12.0%±3.9%)	Towill et al., 1973
	Ophioglossum crotalophoroides	24 h daily	P (36.2%)	Whittier, 2006
	Psilotum complanatumPsilotum nudum	Several months (cultured on nutrient medium, PH= 5.4)	P (52.0% – 98.3%)	Whittier and Braggins, 1994
	Tmesipteris lanceolata,Tmesipteris sigrnatifolia		P
	Tmesipteris elongata		I (0%)
	Tmesipteris tannensis		I (<0.1%)
White light	Adiantum reniforme var. sinense	16 h daily	P (maximum germination rate, 50.1%)	Wu et al., 2010
	Alsophila metteniana	12 h daily	P (68.78%±1.81%)	Du et al., 2009a
	Cheilanthes feei	–	P (98%)	Nondorf et al., 2003
	Lycopodium clavatum	30 min daily or 12 h daily	I (2.2% or 0%)	Whittier, 2008
	Lygodium heterodoxum	12 h daily	P (90%)	Pérez-García et al., 1994
	Ophioglossum crotalophoroides	20 min daily	I (0%)	Whittier, 2006
Green light	Cheilanthes feei	–	P	Nondorf et al., 2003
Yellow light	Alsophila metteniana	12 h daily	P (63.74%±3.06%)	Du et al., 2009a
Blue light	Adiantum capillus-veneris	5 min (followed in darkness for 168 h )	I (0%)	Furuya et al., 1997
	Alsophila metteniana	12 h daily	I (7.51%±0.44%)	Du et al., 2009a
	Cheilanthes feei	–	P (68%)	Nondorf et al., 2003
	Mohria caffrorum	144 h	I (15.7%±3.5%)	Reynolds and Raghavan, 1982
	78 h	I (16%)
	Onoclea sensibilis	5 min	P (51.7%±3.0%)	Towill et al., 1973
	Polypodium aureum	1 d, 3 d, 5 d	I (0%)	Spiess et al., 1977
	Thelypteris kunthii	24 h (followed in drakness for 24 h)	I (0%)	Huckaby and Raghavan, 1981

Tab.1 Effects of different light source on fern spore germination

Environmental factors	Plant species	Treatment condition (concentration, time)	Effects on spore germination	References
R-FR	Adiantum capillus-veneris	R 5 min-FR 5 min (followed in darkness for 168 h)	I (0.4%)	Furuya et al., 1997
	R 38 h-FR 10 min (followed in darkness for 38 h)	P (83.0%±2.2%)
	Asplenium nidus	R 12 h-FR 48 h	I	Raghavan, 1971
	Cheilanthes farinosa	R 30 s-FR 30 s –1 h	N	Raghavan, 1973
	Lycopodium clavatum	R 30 min-FR 30 min daily	P (96.8%)	Whittier, 2008
	Mohria caffrorum	R 12 h-FR 5 min	N (41.1%±2.1%)	Reynolds and Raghavan, 1982
	R 12 h-FR 1 h	N (39.7%±4.4%)
	Ophioglossum crotalophoroides	R 20 min-FR 20 min daily	P (28.4%)	Whittier, 2006
	Thelypteris kunthii	R 8 h-FR 5 min	I (0%)	Huckaby and Raghavan, 1981
FR-R	Adiantum capillus-veneris	FR 5 min-R 5 min (incubated in darkness for 168 h)	P (58.3%)	Furuya et al., 1997
	Asplenium nidus	FR 24–48 h-R 1–2 min	P	Raghavan, 1971
	FR>72 h-R 0–10 min	P
	Lycopodium clavatum	FR 30 min daily-R 30 min daily	I (6.4%)	Whittier, 2008
	Ophioglossum crotalophoroides	FR 20 min daily-R 20 min daily	I (0%)	Whittier, 2006
R-FR-R	Adiantum capillus-veneris	R 5 min-FR 5 min-R 5 min(followed in darkness for 168 h)	P (54.9%)	Furuya et al., 1997
	R 38 h-FR 10 min-R 5 min(followed in darkness for 38 h)	P (72.0%±2.6%)
	Asplenium nidus	R 12 h-FR 48 h-R 10 min	P	Raghavan, 1971
	Thelypteris kunthii	R 8 h-FR 5 min-R 8 h	P (48.2%±4.91%)	Huckaby and Raghavan, 1981
	R 8 h-FR 5 min-R 5 min	I (6.8%±2.69%)
FR-R-FR	Adiantum capillus-veneris	FR 5 min-R 5 min-FR 5 min(followed in darkness for 168 h)	I (0%)	Furuya et al., 1997
R-D-B	Adiantum capillus-veneris	R 5 min-B 5 min (followed in darkness for 168 h)	I (2.6%)	Furuya et al., 1997
	R 38 h-B 5 min (followed in darkness for 38 h)	P (87.3%±1.9%)
	Cheilanthes farinosa	R 30 s-B 30 s	I (0%)	Raghavan, 1973
		R 30 s-D 3 h-B 30 s	SI (50%)
		R 30 s-D 5 h-B 30 s	N
	Mohria caffrorum	R 12 h-B 1 h	I (0.5%±1.6%)	Reynolds and Raghavan, 1982
		R 12 h-B 1–6 h	I
		R 12 h-B 6–72 h	P
	Polypodium aureum	R 1 d-B 1 d; R 1 d-B 3 d; R 1 d-B 5 d	I (11%, 12%, 15%)	Spiess et al., 1977
		R 3 d-B 1 d; R 3 d-B 3 d; R 3 d-B 5 d	SI (49%, 59%, 57%)
		R 5 d-B 1 d; R 5 d-B 3 d; R 5 d-B 5 d	SI (64%, 66%, 76%)
	Thelypteris kunthii	R 8 h-B 24 h	I (17.4%±1.09%)	Huckaby and Raghavan, 1981
B-R	Adiantum capillus-veneris	B 5 min-R 5 min (incubated in darkness for 168 h)	I (2.4%)	Furuya et al., 1997
	Cheilanthes farinosa	B 30 s-R 30 s	I	Raghavan, 1973
		B 30 s-R 1 h	P
		B 30 s-R (high intensity) 1–2 min
	Mohria caffrorum	B 1 h-R 12 h	I (0.3%±1.1%)	Reynolds and Raghavan, 1982
	Polypodium aureum	B 1 d-R 1 d; B 3 d-R 1 d; B 5 d-R 1 d	P (53%; 63%; 63%)	Spiess et al, 1977
		B 1 d-R 3 d; B 3 d-R 3 d; B 5 d-R 3 d	P (68%; 71%; 63%)
		B 1 d-R 5 d; B 3 d-R 5 d; B 5 d-R 5 d	P (52%; 68%; 65%)
	Thelypteris kunthii	B 24 h-R 8 h	I (19.1%±1.47%)	Huckaby and Raghavan, 1981
FR-B	Mohria caffrorum	FR 12 h-B 1 h	I (0%)	Reynolds and Raghavan, 1982
		FR 12 h-B 1–6 h	I
		FR 12 h-B 6–72 h	P
B-FR	Mohria caffrorum	B 1 h, FR 12 h	I (0%)	Reynolds and Raghavan, 1982
R-B-R	Adiantum capillus-veneris	R 5 min-B 5 min-R 5 min(followed in darkness for 168 h)	I (5.4%)	Furuya et al., 1997
	R 38 h-B 5 min-R 5 min(followed in darkness for 38 h)	P (73.9%±3.1%)
	Cheilanthes farinosa	R 30 s-B 30 s-R 1 h	P	Raghavan, 1973
	R 30 s-B 30 s-R (high intensity) 1–2 min	P
R-FR-R-FR	Adiantum capillus-veneris	R 38 h-FR 10 min-R 5 min-FR 10 min(followed in darkness for 38 h)	P (82.6%±2.2%)	Furuya et al., 1997
	Asplenium nidus	R 12 h-FR 48 h-R10 min-FR 8 h	I	Raghavan, 1971
	Thelypteris kunthii	R 8 h-FR 5 min-R 8 h-FR 5 min	P (36.7%±1.83%)	Huckaby and Raghavan, 1981
		R 8 h-FR 5 min-R 5 min-FR 5 min	I (0%)
		R 8 h-FR 5 min-R 8 h-FR 5 min-R 8vh	P (55.2%±4.83%)
		R 8 h-FR 5 min-R 5 min-FR 5 min-R 5 min	I (5.2%±0.92%)
D-R	Asplenium nidus	D 24–72 h-R 1–10 min	I	Raghavan, 1971
Thelypteris kunthii	D 24 h-R 8 h	P (33.0%±3.04%)	Huckaby and Raghavan, 1981
B-R-D-FR	Cheilanthes farinosa	B 30 s-R (high intensity) 2 min-FR 1–2 min	I	Raghavan, 1973
B 30 s-R (high intensity) 2 min-D 5 h-FR 1–2 min	N
B-R-B	Cheilanthes farinosa	B 30 s-R (high intensity) 2 min-B 1–2 min	N	Raghavan, 1973
B 30 s-R (high intensity) 2 min-B 1–2 h	I
R-B-R-D-FR	Cheilanthes farinosa	R 30 s-B 30 s-R (high intensity) 2 min-FR 1–2 min	I	Raghavan, 1973
R 30 s-B 30 s-R (high intensity) 2 min-D 5 h-FR 1–2 min	N
R-B-R-B	Cheilanthes farinosa	R 30 s-B 30 s-R (high intensity) 2 min, B 1–2 min	N	Raghavan, 1973
	R 30 s-B 30 s-R (high intensity) 2 min-B 1–2 h	I
	Adiantum capillus-veneris	R 38 h-B 5 min-R 5 min-B 5 min (incubated in darkness for 38 h)	P (67.9%±3.3%)	Furuya et al., 1997

Tab.2 Effects of multi-light quality interaction on fern spore germination

Fig.2 Effects of light intensity and illumination time on fern spore germination. The red solid lines represent ‘promotion’ and the ‘inhibition’ is shown in green dashed lines. LD₁: Long time of light irradiation (≥24 h); LD₂: Short time irradiation (<24 h); LI-h: High light intensity; LI-l: Low light intensity; LI-s: Leaf shade; LI-w: White light.

Fig.3 Effects of phytohormones on fern spore germination. The red solid lines represent ‘promotion’ and the ‘inhibition’ is shown in green dashed lines, whereas the blue dashed lines indicate no influence on the fern spore germination of current environmental factor. ABA: Abscisic acid; An: Antheridiogen; E: Ethylene (≥1mL·L^-1); E-im: The treatment of ethylene on the immature fern spores; E-m: The treatment of ethylene on the mature spore; GA: Gibberellin acid; GA₄₊₇: A kind of gibberellin acid; JA: Jasmonic acid.

Environmentalfactors	Plant spices	Treatment condition(concentration, time)	Effects on spore germination(germination rate or time)	References
Mineral content in medium and type of Medium	Adiantum raddianum	Knop’s or MS or 1/2 MS	42% or 32% or 48%	Tian et al., 2008
		Knop’s, activated charcoal 0–1.5 g·L^-1	42%–80%
		Knop’s, GA 0–3 mg·L^-1	42%–80%
	Adiantumreniforme var. sinense	1/4 MS, sucrose 0–60 g·L^-1	Maximum germination 61.2%±16.2%	Wu et al., 2010
		1/2 MS, sucrose 0–60 g·L^-1	Maximum germination 41.5%±7.9%
		MS, sucrose 0–60 g·L^-1	Maximum germination 29.8%±6.6%
	Alsophila podophylla	1/2 MS or MS	60.2% or 58.7%	Zhang et al., 2007
	1/10 MS or 1/5 MS	67.5% or 66.9%
	Anemia phyllitidis	Liquid medium	P	Douglas, 1994
	Dryopteris varia	Ms or 1/2 MS or 1/4 MS	33% or 38% or<5%	Ouyang et al., 2008
	Osmunda japonica	1/2 MS or MS	50.3% or 80.2%	Yuan et al., 2002
	1/8 MS	95.3%
	Platycerium bifurcatum	Solid media, 3 d or 6 d	53.5%±5.5% or 67.6%±6.0%	Camloh, 1993
	Liquid media, 3 d or 6 d	50.3%±3.6% or 68.8%±1.4%
	Pteridium aquilinum	Solid media, 14 d	85.2%	Sheffield et al., 2001
	Liquid media, 14 d	60.0%
	Pteris cretica	Knop’s or MS	80.6% or 77.2%	Xu et al., 2005
		1/2 MS	82.3%
		1/5 MS or 1/10 MS	64.7% or 62.5%
	Pteris tripartita Sw	MS	70.66%±1.45%	Baskaran et al., 2012
		1/2 MS	80.00%±1.15%
		1/4 MS or 1/8 MS	59.00%±1.15% or 56.00%±2.51%
	Pteris wallichiana	MS	78.2%±2.1%	Zhang et al., 2008c
		1/2 MS	83.3%±2.3%
		1/5 MS or 1/10 MS	66.7%±2.5% or 62.9%±0.9%
	Pyrrosia lingua	MS	59.5%/9 d	Du et al., 2009b
		1/2 MS or 1/4 MS	82.3%/7d or 82.7%/7d
		1/8 MS or Knop’s	83.2%/6d or 84.1%/6 d
	Schizaea dichotoma	1/4 MS	34%	Cox et al., 2003
	1/2 MS	12%
Sucrose and other contents in culture medium	Athyrium filix-femina	Liquid medium, sucrose 0 M	87.2%	Sheffield et al., 2001
	Liquid medium, sucrose 0.05 M or 0.2 M	91.2% or 94.7%
	Cibotium barometz	leachates of Eupatorium adenophorum(0%, 5%, 10%, 20%, 30%, 40%, 50%)	98% to 18%	Zhang et al., 2008b
	Colysis latiloba	0% Sucrose	15%	Parajuli et al., 2013
		1% Sucrose	86%
		2%–5% Sucrose	75%–54%
	Dryopteris expansa	Liquid medium, sucrose 0 M	57.8%	Sheffield et al., 2001
	Liquid medium, sucrose 0.05 M or 0.2 M	61.1% or 66.0%
	Dryopteris varia	Sucrose 0% or 1% or 2%	37% or 39% or 38%	Ouyang et al., 2008
	Sucrose 3% or 4% or 5%	<5% or <1%
	Macrothelypteris torresiana	leachates of Ageratina adenophora(0%, 5%, 10%, 20%, 30%, 40%, 50%)	nearly 100% to 0%	Zhang et al., 2008a
	Neocheiropteris palmatopedata	leachates of Ageratina adenophora(0%, 5%, 10%, 20%, 30%, 40%, 50%)	nearly 100% to 0%	Zhang et al., 2012
	Osmunda japonica	1/8 MS sucrose 0%	34.8%	Yuan et al., 2002
		1/8 MS, sucrose 2% or 3%	90.2% or 95.1%
		1/8 MS, sucrose 1% or 4% or 5%	80.1% or 83.3% or 64.5%
	Pteridium aquilinum	Liquid medium, sucrose 0 M	30.4%	Sheffield et al., 2001
	Liquid medium, sucrose 0.05 M or 0.2 M	39.3% or 43.3%
	Pteridiumaquilinum var. latiusculum	Sucrose 50000 mg·L^-1 , Vitamin	8.0%	Bao et al., 2000
		Sucrose 50000 mg·L^-1 , Amino acid	9.8%
		Sucrose 50000 mg·L^-1 ,extract of Pteridium aquilinum leaves	20%
		Sucrose 50000 mg·L^-1 , IAA, KN	6.8%
		Sucrose 50000 mg·L^-1 ,(did not add any growth-assisted substance)	6.7%
	Pteris cretica	Sucrose 0% or 1% or 2%	82.3% or 83.0% or 82.6%	Xu et al., 2005
	Sucrose 3% or 4% or 5%	60.5% or 54.7% or 50.1%
	Pyrrosia lingua	Sucrose 0% or 1% or 2%	82.4% or 81.3% or 80.6%	Du et al., 2009b
	Sucrose 3% or 4% or 5%	62.5% or 68.2% or 53.6%
pH	Alsophila metteniana	3.7 or 5.7 or 6.7	65.6%–40%	Du et al., 2009a
		4.7	65.6%
		≥7.7	0%
	Anemia mexicana	pH 5.0–6.5	Maximum germination	Nester and Coolbaugh, 1986
	pH≤4.5	I
	Anemia phyllitidis	pH 5.0–6.5	Maximum germination
	pH≤4.5	I
	Cheilanthes feei	4.5 or 5.5 or 6.5 or 8.5	Nearly 95% or 75% or 55% or 50%	Nondorf et al., 2003
	Cyathea atrovirens	pH 4.0–6.5, 6 d–12 d	78%–93%	Rechenmacher et al., 2010
	pH 7.0, 6 d–12 d	40%–80%
	Drynaria fortunei	7.7	63.3%	Chang et al., 2007
		9.7	0.8%
		3.7	1%
	Gleichenella pectinata	pH 4.0	0.8%±0.4%	Santos et al., 2010
		pH 4.5–5.0	1.2%±0.8%-2.2%±0.8%
		pH 5.5–6.0	0.8%±0.4%-0.5%±0.5%
	Marsilea vestita	pH 7 to 8	78%	Mahlberg and Yarus, 1977
	Ophioglossum palmatum	pH 3.4	14%	Whittier and Moyroud, 1993
		pH 4.0	55%
		pH 4.6–6.2	48%–0%
	Polypodium lepidopteris	pH 4.0–6.7	32%–38%	Viviani and Randi, 2008
Cd²⁺	Ceratopteris thalictroides	0.1 mg·L^-1 or 1.0 mg·L^-1	N	Gupta et al., 1992
2.5 mg·L^-1or 5.0 mg·L^-1	14% or 36%
La³⁺	Dryopteris paleacea	10^-2 M or 10^-4 M or 10^-6 M or 10^-8 M,R 1 min	2.0%±0.7% or 46.4%±2.2%or 58.0%±2.6% or 60.0%±2.4%	Scheuerlein et al., 1989
0 M, R 1 min	60.2%±1.0%
Ca²⁺	Dryopteris paleacea	≤10^-7 M	I (5%)	Scheuerlein et al., 1989
		10^-4 M	Maximum germination
		≥10^-1 M	I

Tab.3 Effect of medium, sucrose and other environmental factors on the germination of fern spores

Fig.4 Effects of temperature and the other environmental factors on fern spore germination. The red solid lines represent ‘promotion’ and the ‘inhibition’ is shown in green dashed lines. CO_2a: CO₂ treatment concentration for the spores of P. piloselloides (219–3360 cm³·m^-3) and O. sensibilis (0–2%); CO_2b: CO₂ treatment concentration for O. sensibilis spores (5%–15%); ID₁: Inoculation density, 50 spores·mL^-1; ID₂: Inoculation density, 3000 spores·cm^-2; T<15: the optimal germination temperature less than 15 °C; Top-20: The maximal germination rate occurred at 20 °C; Top-25: The optimal germination temperature is 25 °C; Top-30: The maximal germination rate occurred at 30 °C.

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