第六讲 海洋底栖藻类生态
近海底栖植物区系
?Benthos 海底生物（包括底栖动、植物）
?近海底栖植物包括：
red algae,brown algae,green algae,blue-green
algae,diatoms,
seagrass,mangroves,marsh grass
?海洋底栖植物生存于陆地与大洋之间的交界地
带，具有高生产力、栖息地类型多变、人类和
动物影响大等特点。
? 海洋底栖植物分布范围：
上限,This region extend on to the land until the effect of
seawater is negligible and the flora indistinguishable from
that of typical freshwater habitats.
下限,The lowermost point seaward must be the point
below which growth of benthic flora ceases due to lack of
light for photosynthesis,and this may be close to land on
steep slope and out on the continental shelf on gentle slope.
底栖植物分布的下限会随着季节和水体混浊度的
变化而变化，这些都和水体中阳光的透射强度变化有
关。
? 底栖植物区系通常都是 light limited（光限制）,由于浮
游藻类、海草甚至陆源植物的遮荫作用，导致水体的
光线透射率降低；例如，如果大型海藻密集生长，其
叶状体（ algal frond）就会形成一个遮蓬，阻挡光线透
射下去，对于小型底栖藻类而言，光就成为限制其生
长的关键因子；
? 海流、波浪运动、陆地径流输入、动物活动等，都会
增加水体混浊度，甚至翻起的泥沙会覆盖藻体，造成
光线不足，限制底栖植物生长；
Estuarine waters carry high concentrations of silt in
suspension and this may restrict light penetration to such
and extent that phytoplankton and benthic production is
severely reduced.
? 底质为底栖植物提供了适宜的附着场所；
? 沉积物及丰富的异养生物（动物、微生物等），使得
底层水体具有丰富的营养盐，一般认为，与外海浮游
植物的营养盐限制相比较，近海底栖植物区系受到营
养盐限制的可能性比较小；
? 浅海水体中，波浪运动、海流和上升流等都能产生搅
动，阻止水体的热力学分层（ except for brief period at
least in the temperate zone），因此，营养盐很少能成
为底栖植物生长的限制因子，与外海相比较，近岸具
有更高的生产力。
? 陆源径流输入和水体中营养盐再循环，使得近海水体
中营养盐含量丰富。
?空间的竞争
In densely populated benthic community,
competition for space is likely to be severe,and
biotic interactions among algae,and between algae
and other groups of organisms like bacteria,fungi,
seagrasses,grazing animals are more complex.
Spores(孢子 ) of seaweeds germinates（萌发）
only when they adheres to substrate,
Crowding lead shading and competition for
nutrients and there is an inverse relationship
between density and individual size.
两种不同的底栖环境
?Intertidal zone 潮间带 in which organisms
are subject at intervals to desiccation (干燥 ),direct
solar heat,rain,snow,freshwater runoff,wind,
frost,etc.
?Subtidal zone 潮下带 which is strictly aquatic
and more uniform,but has nevertheless marked
variables of light intensity and quality.
?The algae growing in intertidal zone must be able
to withstand and recover rapidly from these
changes than subtidal algae,
?Full sunlight severly in hibit photosynthesis in
phytoplanktons,but photosynthesis versus light
intensity curve of benthic diatoms has a broad
optimum,saturating at fairly low light intensities
with only slight inhibition under full sunlight.
两种主要的沉积物基质
?Solid rock 礁石
?Particulate sand or mud 沙、泥
?In general,on rocky shores,the greatest amount of
solid substratum is to be found between tide marks,
with a progressive change through sand to mud in
deep water.
?The stability of substratum is obviously important
for growth of seaweeds,
?The greatest growth of seaweeds occur where there
is a rock surface for attachment.
?The topography（地形） of rocky shore also
affects the type of flora,
On steep rocky surface,silt（泥沙） doesn’t
accumulate and only epilithic （长在石头上的）
flora grow,But a change in the flora occur where
sufficient silt of sand collect on flat rocky surface.
Some epilithic algae,Laminaria 海带
Porphyra 紫菜
? The moving pebbles(鹅卵石 ) or gravel(砂砾 ) can not
maintain an epilithic flora,yet similar sized pebbled in a
protected habitat are colonized by genera such as
Enteromorpha(浒苔 ),Ectocarpus（水云） of even Fucus
（墨角藻）,
? In areas of sand movement,the species diversity is low
and the species tend to be tough and wiry,
? Most of the seaweeds found on sandy shore are torn loose
from their holdfast（固着器） and washed on to shore.
第一节 微型底栖藻类
?Types of benthic microalgae 底栖微藻的类群
1) Blue-green algae 蓝绿藻
2) Diatom 硅藻
3) Dinoflagellates 甲藻
1） Blue-green algae
?In marine habitats,blue-green algae are relatively
inconspicuous as compared to other algae,The
growth of blue-green algae often gives a black
appearance to the substratum.
?Blue-green algae live in all parts of the lighted
marine benthic environment and may be dredged
from water depth more than 30m under very dim
illumination.
?Blue-green algae on mud-flats make a significant
contribution to the nitrogen balance and also exert
a profound stabilizing influence on the substratum.
蓝绿藻的固氮作用，将氮气直接转化为可为植
物直接利用的氮盐。
一些常见的近海蓝绿藻（ Fig.）
2) Diatom
?The most ubiquitous （普遍存在的） of all
benthic marine algae are diatoms.
?They are either attached to or lie loose on the
rocky,sandy or muddy surface.
? The physical nature of the shore has a profound influence
on the flora,海滩的物理环境对植物区系有重要影响
On coarse sandy beaches,motile diatom species seems
to be absent and only those like Achnanthes（曲壳藻）
and Cocconeis（卵形藻） which are attached to sand
particles are mostly found,whereas on fine sand or on
silty mud,the species of Nitzschia（菱形藻）,Navicula
（舟形藻）,Pleurosigma（斜纹藻）,Amphora（双眉
藻）,Caloneis（美壁藻）,Diploneis（双壁藻）,etc,
which lie loose on the surface often from a brown
coloration during stable period.
? 底栖硅藻的垂直移动
? Benthic diatom flora lying loose on the surface shows
diurnal vertical movement,they disappear from the
surface when covered by tide and reappear when exposed
during low tide provided that light is sufficient,
? This is probably a mechanism to avoid wash-out and
abrasion(磨损 ) damage in the disturbed surface layers at
high tide,however many benthic diatoms will be put into
suspension by incoming tide.
?In sublittoral(近海滩的 ) zone,the number of
diatoms fall presumably because of decrease in
light intensity with depth.
3） Dinoflagellates
?Patches of dinoflagellates occur on sandy beaches
and cause a greenish brown discoloration(变色、
污点 ) of the sand.
?The discoloration of sand disappears shortly before
it becomes covered by the rising tide and reappears
when the sand is again exposed.
?All of these sand-inhabiting forms are flat and
generally very mobile.
第二节 大型底栖藻类
?What are seaweeds?
Seaweeds are classified as algae,photosynthetic
organisms which,in contrast to the plants growing
on land,are non-flowering and do not have roots,
leafy shoots or sophisticated tissues for
transporting water,sugars and nutrients.
? 大型绿藻、褐藻、红藻通过固着器或者假根固着生在
在岩石基质上；
? They grow extensively extending vertically from the
upper limit to which wave splash reaches,downward
through the intertidal to the subtidal up to the lower limit
of photic zone.分布的上限为潮流与波浪能到达的地方，
下限为真光层的下限；
? Mixed with seaweeds are also present large number of
microscopic benthic blue-green algae and diatoms.
?Seaweeds are mostly confined to permanent rocky
shores and are usually absent of less frequent from
shores of shifting sand,gravel and cobbles(鹅卵
石 ),because they can not ordinarily survive the
movement of current and surge without secure
attachment to substratum.
? Holding fast 固着
Roots are not needed for seaweeds since nutrients such
as nitrogen,phosphorus and trace elements are dissolved
in the seawater,The nutrients can be taken up and
exchanged by diffusion and active transport directly
across all the surfaces of the seaweed (carbon dioxide and
oxygen are also assimilated and released in this way),
Therefore it is only the mechanical features of a root
system that would be beneficial for the seaweeds,holding
them steady no matter how turbulent the water movement
is.
?Some seaweeds,such as the conchocelis phase(壳
孢子世代 ) of Porphyra,obtain a foothold using a
more aggressive approach,They secrete acid thich
dissolves the calcium carbonate(碳酸钙 ) of shells
and coral skeletons so that fine rhizoids and
filaments can grow,The calcium and bicarbonate
that are released by the acidification of the calcium
carbonate structures also provide a localized
source of nutrients for the seaweed.
?The quest for light
In cloudy coastal waters full of plankton and
suspended particles washed in from the land and
brought down by rivers,light may not penetrate
further than a few meters,There are several
structural features,common to many species of
seaweed,that help to address this problem.
? (1) the stipe (柄 )
The stipe of a seaweed can be very long indeed,The
purpose is to support the bulk(主要部分 ) of the
photosynthetic tissue at the surface of the water where
incident light is at its maximum intensity.
例如,Nereocystis luetkeana（腔囊藻）的柄长可达
30m,可支撑超过 100张叶片，每张叶片几米长，长柄
可使得叶片尽量获取更多的光能进行光合作用。
?(2) Buoyancy aids 增加浮力的构造 — 气囊
Large gas-filled bladder ensures that the blades
are floating as close to the surface of the water as
possible.
例如,Macrocystis pyrifera (巨藻 ) 的每张叶
片上都有气囊，有助于整棵植物体能悬浮在水
体中，尽量获得更多的光线。
? In spite of superficial resemblance of kelp holdfasts to the
roots of vascular plants,there is no evidence that they are
supplying nutrients to other parts of the plant,The whole
thallus (叶状体 ) of seaweeds performs photosynthetic and
absorptive functions.
? Large,long-lived thallus of seaweeds enable them to store
nutrients,which extend their growing periods and
represent a survival strategy not available to microscopic
algae.
Type of seaweeds
?1) Green seaweeds
?2) Brown seaweeds
?3) Red seaweeds
1) Green seaweeds
?Green algae are relatively of minor occurrence in
sea as compared with their overwhelming
dominance in freshwaters,and unlike in
freshwaters,marine green algae are primarily
macroscopic forms.
2) Brown seaweeds
?They occur in bulk in nearly all temperate and
high-latitude waters,where they dominant the
vegetation of coastal waters,but they are much
smaller in number and size in tropical coastal
water.
?Sargassum(马尾藻 ) is a classic example of an
abundant free-floating brown alga in Sargasso sea.
?Brown algae are in large part macroscopic algae
with no unicellular,colonial or unbranched
filamentous form.
?The smallest form such as Ectocarpus(水云 ) is
consisting of branched uniseriate（单列的）
filaments,while the largest members of Fucales
（墨角藻） and Laminariales（海带） exhibit a
much bigger size and an elaboration of forms.
3） Red seaweeds
?Red algae occupy the entire range of habitats from
highest intertidal to lowermost limits of light
penetration,but majority of them prefer deep water.
?Almost all of them grow attached to rocks or other
algae.
? Although red seaweeds occur in all latitudes,there is a
marked shift in their proportional abundance as one moves
from high to low latitudes.
? There are only few red algal species occurring in north
polar and subpolar regions(where brown algae
predominates),but they far outnumber the brown and
green seaweeds in tropical and temperate regions.
? Large fleshy red algal species usually occur mostly in
cool-temperate areas,whereas massive calcareous(石灰质
的 ) forms and small filamentous forms occur mostly in
tropical seas.
近海底栖植物区系
?Benthos 海底生物（包括底栖动、植物）
?近海底栖植物包括：
red algae,brown algae,green algae,blue-green
algae,diatoms,
seagrass,mangroves,marsh grass
?海洋底栖植物生存于陆地与大洋之间的交界地
带，具有高生产力、栖息地类型多变、人类和
动物影响大等特点。
? 海洋底栖植物分布范围：
上限,This region extend on to the land until the effect of
seawater is negligible and the flora indistinguishable from
that of typical freshwater habitats.
下限,The lowermost point seaward must be the point
below which growth of benthic flora ceases due to lack of
light for photosynthesis,and this may be close to land on
steep slope and out on the continental shelf on gentle slope.
底栖植物分布的下限会随着季节和水体混浊度的
变化而变化，这些都和水体中阳光的透射强度变化有
关。
? 底栖植物区系通常都是 light limited（光限制）,由于浮
游藻类、海草甚至陆源植物的遮荫作用，导致水体的
光线透射率降低；例如，如果大型海藻密集生长，其
叶状体（ algal frond）就会形成一个遮蓬，阻挡光线透
射下去，对于小型底栖藻类而言，光就成为限制其生
长的关键因子；
? 海流、波浪运动、陆地径流输入、动物活动等，都会
增加水体混浊度，甚至翻起的泥沙会覆盖藻体，造成
光线不足，限制底栖植物生长；
Estuarine waters carry high concentrations of silt in
suspension and this may restrict light penetration to such
and extent that phytoplankton and benthic production is
severely reduced.
? 底质为底栖植物提供了适宜的附着场所；
? 沉积物及丰富的异养生物（动物、微生物等），使得
底层水体具有丰富的营养盐，一般认为，与外海浮游
植物的营养盐限制相比较，近海底栖植物区系受到营
养盐限制的可能性比较小；
? 浅海水体中，波浪运动、海流和上升流等都能产生搅
动，阻止水体的热力学分层（ except for brief period at
least in the temperate zone），因此，营养盐很少能成
为底栖植物生长的限制因子，与外海相比较，近岸具
有更高的生产力。
? 陆源径流输入和水体中营养盐再循环，使得近海水体
中营养盐含量丰富。
?空间的竞争
In densely populated benthic community,
competition for space is likely to be severe,and
biotic interactions among algae,and between algae
and other groups of organisms like bacteria,fungi,
seagrasses,grazing animals are more complex.
Spores(孢子 ) of seaweeds germinates（萌发）
only when they adheres to substrate,
Crowding lead shading and competition for
nutrients and there is an inverse relationship
between density and individual size.
两种不同的底栖环境
?Intertidal zone 潮间带 in which organisms
are subject at intervals to desiccation (干燥 ),direct
solar heat,rain,snow,freshwater runoff,wind,
frost,etc.
?Subtidal zone 潮下带 which is strictly aquatic
and more uniform,but has nevertheless marked
variables of light intensity and quality.
?The algae growing in intertidal zone must be able
to withstand and recover rapidly from these
changes than subtidal algae,
?Full sunlight severly in hibit photosynthesis in
phytoplanktons,but photosynthesis versus light
intensity curve of benthic diatoms has a broad
optimum,saturating at fairly low light intensities
with only slight inhibition under full sunlight.
两种主要的沉积物基质
?Solid rock 礁石
?Particulate sand or mud 沙、泥
?In general,on rocky shores,the greatest amount of
solid substratum is to be found between tide marks,
with a progressive change through sand to mud in
deep water.
?The stability of substratum is obviously important
for growth of seaweeds,
?The greatest growth of seaweeds occur where there
is a rock surface for attachment.
?The topography（地形） of rocky shore also
affects the type of flora,
On steep rocky surface,silt（泥沙） doesn’t
accumulate and only epilithic （长在石头上的）
flora grow,But a change in the flora occur where
sufficient silt of sand collect on flat rocky surface.
Some epilithic algae,Laminaria 海带
Porphyra 紫菜
? The moving pebbles(鹅卵石 ) or gravel(砂砾 ) can not
maintain an epilithic flora,yet similar sized pebbled in a
protected habitat are colonized by genera such as
Enteromorpha(浒苔 ),Ectocarpus（水云） of even Fucus
（墨角藻）,
? In areas of sand movement,the species diversity is low
and the species tend to be tough and wiry,
? Most of the seaweeds found on sandy shore are torn loose
from their holdfast（固着器） and washed on to shore.
第一节 微型底栖藻类
?Types of benthic microalgae 底栖微藻的类群
1) Blue-green algae 蓝绿藻
2) Diatom 硅藻
3) Dinoflagellates 甲藻
1） Blue-green algae
?In marine habitats,blue-green algae are relatively
inconspicuous as compared to other algae,The
growth of blue-green algae often gives a black
appearance to the substratum.
?Blue-green algae live in all parts of the lighted
marine benthic environment and may be dredged
from water depth more than 30m under very dim
illumination.
?Blue-green algae on mud-flats make a significant
contribution to the nitrogen balance and also exert
a profound stabilizing influence on the substratum.
蓝绿藻的固氮作用，将氮气直接转化为可为植
物直接利用的氮盐。
一些常见的近海蓝绿藻（ Fig.）
2) Diatom
?The most ubiquitous （普遍存在的） of all
benthic marine algae are diatoms.
?They are either attached to or lie loose on the
rocky,sandy or muddy surface.
? The physical nature of the shore has a profound influence
on the flora,海滩的物理环境对植物区系有重要影响
On coarse sandy beaches,motile diatom species seems
to be absent and only those like Achnanthes（曲壳藻）
and Cocconeis（卵形藻） which are attached to sand
particles are mostly found,whereas on fine sand or on
silty mud,the species of Nitzschia（菱形藻）,Navicula
（舟形藻）,Pleurosigma（斜纹藻）,Amphora（双眉
藻）,Caloneis（美壁藻）,Diploneis（双壁藻）,etc,
which lie loose on the surface often from a brown
coloration during stable period.
? 底栖硅藻的垂直移动
? Benthic diatom flora lying loose on the surface shows
diurnal vertical movement,they disappear from the
surface when covered by tide and reappear when exposed
during low tide provided that light is sufficient,
? This is probably a mechanism to avoid wash-out and
abrasion(磨损 ) damage in the disturbed surface layers at
high tide,however many benthic diatoms will be put into
suspension by incoming tide.
?In sublittoral(近海滩的 ) zone,the number of
diatoms fall presumably because of decrease in
light intensity with depth.
3） Dinoflagellates
?Patches of dinoflagellates occur on sandy beaches
and cause a greenish brown discoloration(变色、
污点 ) of the sand.
?The discoloration of sand disappears shortly before
it becomes covered by the rising tide and reappears
when the sand is again exposed.
?All of these sand-inhabiting forms are flat and
generally very mobile.
第二节 大型底栖藻类
?What are seaweeds?
Seaweeds are classified as algae,photosynthetic
organisms which,in contrast to the plants growing
on land,are non-flowering and do not have roots,
leafy shoots or sophisticated tissues for
transporting water,sugars and nutrients.
? 大型绿藻、褐藻、红藻通过固着器或者假根固着生在
在岩石基质上；
? They grow extensively extending vertically from the
upper limit to which wave splash reaches,downward
through the intertidal to the subtidal up to the lower limit
of photic zone.分布的上限为潮流与波浪能到达的地方，
下限为真光层的下限；
? Mixed with seaweeds are also present large number of
microscopic benthic blue-green algae and diatoms.
?Seaweeds are mostly confined to permanent rocky
shores and are usually absent of less frequent from
shores of shifting sand,gravel and cobbles(鹅卵
石 ),because they can not ordinarily survive the
movement of current and surge without secure
attachment to substratum.
? Holding fast 固着
Roots are not needed for seaweeds since nutrients such
as nitrogen,phosphorus and trace elements are dissolved
in the seawater,The nutrients can be taken up and
exchanged by diffusion and active transport directly
across all the surfaces of the seaweed (carbon dioxide and
oxygen are also assimilated and released in this way),
Therefore it is only the mechanical features of a root
system that would be beneficial for the seaweeds,holding
them steady no matter how turbulent the water movement
is.
?Some seaweeds,such as the conchocelis phase(壳
孢子世代 ) of Porphyra,obtain a foothold using a
more aggressive approach,They secrete acid thich
dissolves the calcium carbonate(碳酸钙 ) of shells
and coral skeletons so that fine rhizoids and
filaments can grow,The calcium and bicarbonate
that are released by the acidification of the calcium
carbonate structures also provide a localized
source of nutrients for the seaweed.
?The quest for light
In cloudy coastal waters full of plankton and
suspended particles washed in from the land and
brought down by rivers,light may not penetrate
further than a few meters,There are several
structural features,common to many species of
seaweed,that help to address this problem.
? (1) the stipe (柄 )
The stipe of a seaweed can be very long indeed,The
purpose is to support the bulk(主要部分 ) of the
photosynthetic tissue at the surface of the water where
incident light is at its maximum intensity.
例如,Nereocystis luetkeana（腔囊藻）的柄长可达
30m,可支撑超过 100张叶片，每张叶片几米长，长柄
可使得叶片尽量获取更多的光能进行光合作用。
?(2) Buoyancy aids 增加浮力的构造 — 气囊
Large gas-filled bladder ensures that the blades
are floating as close to the surface of the water as
possible.
例如,Macrocystis pyrifera (巨藻 ) 的每张叶
片上都有气囊，有助于整棵植物体能悬浮在水
体中，尽量获得更多的光线。
? In spite of superficial resemblance of kelp holdfasts to the
roots of vascular plants,there is no evidence that they are
supplying nutrients to other parts of the plant,The whole
thallus (叶状体 ) of seaweeds performs photosynthetic and
absorptive functions.
? Large,long-lived thallus of seaweeds enable them to store
nutrients,which extend their growing periods and
represent a survival strategy not available to microscopic
algae.
Type of seaweeds
?1) Green seaweeds
?2) Brown seaweeds
?3) Red seaweeds
1) Green seaweeds
?Green algae are relatively of minor occurrence in
sea as compared with their overwhelming
dominance in freshwaters,and unlike in
freshwaters,marine green algae are primarily
macroscopic forms.
2) Brown seaweeds
?They occur in bulk in nearly all temperate and
high-latitude waters,where they dominant the
vegetation of coastal waters,but they are much
smaller in number and size in tropical coastal
water.
?Sargassum(马尾藻 ) is a classic example of an
abundant free-floating brown alga in Sargasso sea.
?Brown algae are in large part macroscopic algae
with no unicellular,colonial or unbranched
filamentous form.
?The smallest form such as Ectocarpus(水云 ) is
consisting of branched uniseriate（单列的）
filaments,while the largest members of Fucales
（墨角藻） and Laminariales（海带） exhibit a
much bigger size and an elaboration of forms.
3） Red seaweeds
?Red algae occupy the entire range of habitats from
highest intertidal to lowermost limits of light
penetration,but majority of them prefer deep water.
?Almost all of them grow attached to rocks or other
algae.
? Although red seaweeds occur in all latitudes,there is a
marked shift in their proportional abundance as one moves
from high to low latitudes.
? There are only few red algal species occurring in north
polar and subpolar regions(where brown algae
predominates),but they far outnumber the brown and
green seaweeds in tropical and temperate regions.
? Large fleshy red algal species usually occur mostly in
cool-temperate areas,whereas massive calcareous(石灰质
的 ) forms and small filamentous forms occur mostly in
tropical seas.
