跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

富宏牛肉麵適合第一次吃嗎？

走完這 10 家，

你可能會發現一件事胖老闆誠意肉粥好吃嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，頃刻間綠豆沙牛奶專賣店真的推薦嗎？

等你有一天，再回到這座城市。藍家割包有名是真的嗎？

如果你是第一次來臺北，頃刻間綠豆沙牛奶專賣店怎麼點比較好？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，胡記米粉湯會不會太鹹？

也不用為了排行而奔波，頃刻間綠豆沙牛奶專賣店男生會吃得飽嗎？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，胖老闆誠意肉粥真的推薦嗎？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，圓環邊蚵仔煎名過其實嗎？

然後在心裡對自己說一句：胡記米粉湯好吃嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，胖老闆誠意肉粥適合第一次吃嗎？

會一直在這些地方，

等你再回來。

Researchers discovered that Earth’s biodiversity 800 million years ago was richer than previously thought, identifying ancient lineages of organisms that diversified well before the Cambrian explosion. Their findings challenge long-held views about the Neoproterozoic era and highlight the adaptability of early life forms. A recent study suggests that by the Neoproterozoic period, distinct lineages of amoebae, as well as the ancestors of plants, algae, and animals, had already emerged and managed to survive the two global glaciations that covered the planet. Approximately 800 million years ago (mya), long before the formation of the supercontinent Pangea, Earth’s biodiversity was more varied than previously thought. Brazilian researchers, through the reconstruction of the evolutionary tree of life from ancient amoebas and the ancestors of algae, fungi, plants, and animals, have proposed a scenario where multiple distinct lineages of species coexisted during this era. Their findings are detailed in an article published in the Proceedings of the National Academy of Sciences of the United States of America (PNAS). According to the literature, several lineages of eukaryotes that first emerged 1.5 billion years ago diversified and established themselves during the Neoproterozoic oxygenation event (850-540 mya), when oxygen levels in the atmosphere and oceans rose significantly owing to changes in the planet’s geochemistry. Eukaryotes are organisms consisting of one or more cells in which the DNA is contained within a distinct nucleus (all life on Earth except bacteria and archaea). The study conducted by the researchers focused on the origin and divergence times of amoebozoans, showing that many of these organisms, as well as ancestors of plants, algae, fungi, and animals, survived even the two glaciations of the Cryogenian period (790-635 mya, the middle Neoproterozoic Era, preceded by the Tonian and followed by the Ediacaran). According to the snowball earth hypothesis, polar ice extended to cover the entire planet for some 100 million years during this period. Challenging Classical Theories “The classical paradigm for the Neoproterozoic was that there was practically no life on the planet apart from one or two species of bacteria and protists. In the last 15 years, however, fossils of unicellular, eukaryotic, and heterotrophic organisms have been identified at various different locations around the world. These fossils date from about 800 mya [and are termed Tonian]. All this joined our study, which reconstituted the tree of life and used maximum likelihood estimation to identify several well-established Tonian lineages of ancestors of amoebae, animals, fungi, and plants. This radically changes the paradigm for the manner in which the diversification of life occurred on our planet,” Daniel Lahr, last author of the article and a professor at the University of São Paulo’s Institute of Biosciences (IB-USP), told Agência FAPESP. Left, a fossilized thecamoebian believed to have lived 720-635 mya. Right, a specimen from a group of modern amoebozoan testate amoebae. Credit: Luana Morais and João Alcino In other words, the study dates the mass diversification of life on Earth to some 260 million years earlier than the paradigm, well before the Cambrian explosion (the emergence of new multicellular organisms at the beginning of the Cambrian period between 541 and 530 mya). During this period, the Earth was inhabited mostly by marine invertebrates such as trilobites, brachiopods, and graptolites, and had a warm and humid climate with no evidence of glaciers. “The eukaryotes remained highly diverse despite all the climate changes that occurred during the Neoproterozoic, displaying greater adaptability than expected. This is important because our reconstitution of the phylogenetic tree also serves as a basis for paleoclimate reconstruction research,” Lahr explained. “A curious aspect is that Arcellinid amoebae then lived in saltwater, whereas now they all live in freshwater. This kind of change is frequently observed in the course of evolution since time began, but it happened to all lineages in the case of these amoebae, showing once again how adaptable these organisms were.” The researchers deployed innovative techniques to reconstitute the phylogenetic (evolutionary) tree of the genus Thecamoeba, which belongs to the order Arcellinida, and on this basis to rebuild the tree of life, starting with the prime ancestors of plants, fungi, algae, and animals. “The thecamoebians were the foundation for this reconstitution. From there on up, we were able to visualize other organisms that preceded other groups and must also have been present and diversified in the period in question some 800 mya,” Lahr said. Reconstructing the Phylogenetic Tree Previous research had revealed eight new ancestral lineages of Thecamoebae, the largest group in Amoebozoa, a clade (a common ancestor and all its descendants) of protozoans with pseudopodia shaped like tubes or flat lobes used for locomotion and feeding. Thecamoebians are known as testate amoebae because of their hard carapace. “With the aid of mathematical probability modeling, we were able to determine the morphology of ancestral thecamoebians [from genetic data for species now alive on Earth] and compare it with the fossil morphology. In this study we identified ancestral thecamoebian lineages and the augmented diversification of these organisms in the Neoproterozoic,” Lahr said. In the study described in PNAS, the scientists advanced their understanding of life on the planet 800 mya by using these thecamoebian lineages as points in calibrating the phylogenetic tree for plants, algae, fungi, animals and their ancestors. FAPESP supported the work via a regular research grant and a PhD scholarship. “In developing this extension of the tree of life, we made some interesting discoveries about a period in the planet’s history that has always been obscure. By calibrating the tree in accordance with the phylogenetic study of Thecamoebae, we were able to double the amount of information about eukaryotes in the Neoproterozoic. Our analysis of the data showed that a great diversity of lineages began to emerge in the period, one of which was animals and another fungi, possibly alongside plants,” Lahr said. One of the innovative techniques used in the study, he explained, was single-cell transcriptomics, whereby the entire transcriptome of a single cell or unicellular organism can be sequenced. The transcriptome is the set of all RNA transcripts, including coding and non-coding, in an individual or a population of cells. Transcription, the first step in gene expression, involves copying a gene’s DNA sequence to make an RNA molecule. Transcriptome sequencing enables researchers to trace the evolutionary process in reverse, identifying species that lived in the past. “Before this technique was invented, it was possible to obtain only transcriptomes of single-cell organisms living in culture, meaning less than 1% of the full diversity of microorganisms. Thanks to this innovation, we were able to structure the phylogeny of Thecamoebae as a whole. It’s a highly diverse group and is interesting because it illuminates these periods in Earth’s history that have fossil records with which we can make comparisons. Aside from this, in the tree of life, the amoebae are closer to the animals than to the plants, and this discovery enabled us to make important calibrations,” Lahr said. Reference: “Amoebozoan testate amoebae illuminate the diversity of heterotrophs and the complexity of ecosystems throughout geological time” by Alfredo L. Porfirio-Sousa, Alexander K. Tice, Luana Morais, Giulia M. Ribeiro, Quentin Blandenier, Kenneth Dumack, Yana Eglit, Nicholas W. Fry, Maria Beatriz Gomes E Souza, Tristan C. Henderson, Felicity Kleitz-Singleton, David Singer, Matthew W. Brown and Daniel J. G. Lahr, 16 July 2024, Proceedings of the National Academy of Sciences. DOI: 10.1073/pnas.2319628121 The study was funded by the São Paulo Research Foundation.

The salamander species studied by the Yun group: a red spotted newt Notophthalmus viridescens. By studying salamanders, which have remarkable regenerative abilities, researchers discovered that the presence of senescent cells speeds up the limb regeneration process. The cells secrete factors that signal mature muscle fibers to dedifferentiate into muscle progenitor cells, enhancing regeneration. This finding could help researchers understand why humans have limited regenerative abilities and potentially develop new treatments for age-related diseases. Credit: Maximina Yun Scientists show that so-called senescent cells, i.e., cells that have permanently stopped dividing, boost production of new muscle cells to enhance regeneration of lost limbs in salamanders. Senescent cells, linked to aging and disease, may also have regenerative properties. Studying salamanders, researchers found senescent cells accelerate limb regeneration by signaling muscle fibers to dedifferentiate, potentially leading to new treatments for age-related diseases. Senescent cells are cells that have permanently stopped dividing in response to cellular stress but have not died. As organisms age, the number of senescent cells in the body increases. This accumulation is currently considered one of the hallmarks of aging and has been linked to a variety of diseases, including cancer. However, the true nature of these cells may be more complex and context-dependent. A growing body of evidence suggests that senescent cells may also have beneficial effects, such as wound healing or preventing tissue scarring. “A few years ago, our group found that senescent cells were present at key stages of salamander limb regeneration. Interestingly, other groups subsequently found these cells in other regeneration contexts, including in mammals. We, therefore, wanted to find out whether these cells contribute in any way to regeneration itself,” explains Dr. Maximina Yun, research group leader at the Center for Regenerative Therapies Dresden (CRTD) and the Cluster of Excellence Physics of Life (PoL) at TU Dresden and the Max Planck Institute of Molecular Cell Biology and Genetics (MPI-CBG). Senescent Cells Promote Regeneration The researchers in Yun’s group study salamanders. These animals have unique regeneration abilities and are able to re-grow many organs of their bodies, including lost limbs. “Salamander limb regeneration is a fascinating process. In a matter of weeks, they re-grow a fully functional limb,” explains Dr. Yun. To check if the presence of senescent cells influences the limb regeneration process, researchers in the Yun group found a way to modulate the number of senescent cells in the wound. The team observed that the presence of senescent cells enhanced the regeneration process. “When more senescent cells were present in the wound, the animals developed a larger regeneration bud, or – as we call it – blastema. This is a collection of cells that are going to form all the needed tissues in the new limb. The larger the blastema, the more cells are there to regrow the limb and the quicker the regeneration process. The presence of senescent cells seemed to ‘fuel’ the regeneration process,” Dr. Yun says. “Zombie” Signaling Promotes New Muscle Cells Looking more closely at the blastema with and without the influence of the senescent cells, the Yun team uncovered a new mechanism that enhances the regeneration process and found that the presence of senescent cells increased the number of regenerating muscle cells. They showed that senescent cells secrete factors that stimulate nearby muscle tissue to take a developmental step back and produce new muscle. “Our results show that senescent cells use cell-cell communication to influence the regeneration process. They secrete molecules that signal to mature muscle fibers to dedifferentiate into muscle progenitor cells. These cells can multiply themselves as well as differentiate into new muscle cells, thereby enhancing the regeneration process. This signaling appears to be an important part of promoting regeneration,” says Dr. Yun. For now, the group focused on muscle, one of the most important tissues in the regenerating limb. However, the team is already investigating whether senescent cell signaling also contributes to the regeneration of other tissues. Lessons From the Salamanders Yun’s group is working with salamanders to study regeneration and aging processes. “Salamanders are one of the few animal species that seem to defy the natural aging process. They do not develop typical signs of aging and do not accumulate age-related diseases such as cancer. They also have extraordinary healing abilities,” says Dr. Yun. The animals can regenerate almost any organ in their body. Studying salamanders is helping Dr. Yun and her colleagues at the CRTD understand the principles of the regeneration process and, in the long run, may help solve the puzzle of why humans have very limited regenerative abilities. Reference: “Senescent cells enhance newt limb regeneration by promoting muscle dedifferentiation” by Hannah E. Walters, Konstantin E. Troyanovskiy, Alwin M. Graf and Maximina H. Yun, 6 April 2023, Aging Cell. DOI: 10.1111/acel.13826

Fossil of the early sail-backed synapsid Dimetrodon, from 290 million years ago, one of the species investigated in the study. Credit: Christina Byrd. Museum of Comparative Zoology, © President and Fellows of Harvard College A recent study by Harvard reveals that the evolutionary transition to an upright posture in mammals was complex and occurred later than assumed, based on advanced biomechanical modeling and fossil analysis. Mammals, including humans, stand out with their uniquely upright posture, a key trait that has driven their remarkable evolutionary success. However, the earliest known ancestors of modern mammals resembled reptiles more closely, with limbs extending out to the sides in a sprawled posture. The Evolutionary Shift in Posture The transition from a sprawled stance, similar to that of lizards, to the upright posture of modern mammals such as humans, dogs, and horses, marked a critical turning point in evolution. This change involved a major reorganization of limb anatomy and function in synapsids—the group that includes both mammals and their non-mammalian ancestors—eventually leading to the therian mammals (marsupials and placentals) we know today. Despite over a century of research, the precise details of how, why, and when this evolutionary shift occurred have remained elusive. Land animals exhibit a continuum of limb postures – ranging from ‘sprawled’, with the limbs held out to the side of the body, like lizards, to ‘upright’ or ‘erect’, with the limbs held beneath the body and close to the animal’s midline, like dogs, cats, and horses. Upright posture is characteristic of most modern mammals, but when did this key trait evolve? Credit: Peter Bishop New Insights from Harvard Researchers In a new study published in Science Advances, Harvard researchers provide new insights into this mystery, revealing the shift from a sprawled to upright posture in mammals was anything but straightforward. Using cutting-edge methods that blend fossil data with advanced biomechanical modeling, the researchers found that this transition was surprisingly complex and nonlinear, and occurred much later than previously believed. The study involved digitizing the fossil skeletons of extinct synapsids, creating digital biomechanical models of the musculoskeletal system of the hindlimb, and using these models to compute the limb’s ability to apply force on the ground in different directions. The result is a three-dimensional ‘feasible force space’, which describes what the limb is capable of achieving during locomotion. Credit: Peter Bishop Biomechanical Analysis and Fossil Studies Lead author Dr. Peter Bishop, a postdoctoral fellow, and senior author Professor Stephanie Pierce, both in the Department of Organismic and Evolutionary Biology at Harvard, began by examining the biomechanics of five modern species that represent the full spectrum of limb postures, including a tegu lizard (sprawled), an alligator (semi-upright), and a greyhound (upright). “By first studying these modern species, we greatly improved our understanding of how an animal’s anatomy relates to the way it stands and moves,” said Bishop. “We could then put it into an evolutionary context of how posture and gait actually changed from early synapsids through to modern mammals.” Evolutionary interrelationships of the modern (black silhouettes) and extinct (gray silhouettes) species investigated. The study revealed a complex history of posture evolution in synapsids, and that a fully ‘upright’ posture typical of modern placentals and marsupials was late to evolve. Credit: Peter Bishop Advanced Computational Models and Their Implications The researchers extended their analysis to eight exemplar fossil species from four continents spanning 300 million years of evolution. The species ranged from the 35g proto-mammal Megazostrodon to the 88kg Ophiacodon and included iconic animals like the sail-backed Dimetrodon and the saber-toothed predator Lycaenops. Using principles from physics and engineering, Bishop and Pierce built digital biomechanical models of how the muscles and bones are attached to each other. These models allowed them to generate simulations that determined how much force the hindlimbs (back legs) could apply on the ground. “The amount of force that a limb can apply to the ground is a critical determinant of locomotor performance in animals,” said Bishop. “If you cannot produce sufficient force in a given direction when it’s needed, you won’t be able to run as fast, turn as quickly, or worse still, you could well fall over.” The computer simulations produced a three-dimensional “feasible force space” that captures a limb’s overall functional performance. “Computing feasible force spaces implicitly accounts for all the interactions that can occur between muscles, joints and bones throughout a limb,” said Pierce. “This gives us a clearer view of the bigger picture, a more holistic view of limb function and locomotion and how it evolved over hundreds of millions of years.” While the concept of a feasible force space (developed by biomedical engineers) has been around since the 1990s, this study is the first to apply it to the fossil record to understand how extinct animals once moved. The authors packaged the simulations into new “fossil-friendly” computational tools that can aid other paleontologists in exploring their own questions. These tools could also help engineers design better bio-inspired robots that can navigate complex or unstable terrain. Revisiting Extinct Species’ Locomotor Performance The study revealed several important ‘signals’ of locomotion, including that the overall force-generating ability in the modern species was maximal around the postures that each species used in their daily behavior. Importantly, this meant that Bishop and Pierce could be confident that the results obtained for the extinct species genuinely reflected how they stood and moved when alive. Fossil of the mammal-like cynodont Massetognathus, from 242 million years ago, one of the species investigated in the study. Credit: Peter Bishop. Museum of Comparative Zoology, © President and Fellows of Harvard College Implications for the Evolution of Mammalian Posture After analyzing the extinct species, the researchers discovered that locomotor performance peaked and dipped over millions of years, rather than progressing in a simple, linear fashion from sprawling to upright. Some extinct species also appeared to be more flexible—able to shift back and forth between more sprawled or more upright postures, like modern alligators and crocodiles do. While others showed a strong reversal towards more sprawled postures before mammals evolved. Paired with the study’s other results, this indicated that the traits associated with upright posture in today’s mammals evolved much later than previously thought, most likely close to the common ancestor of therians. These findings also help reconcile several unresolved problems in the fossil record. For example, it explains the persistence of asymmetric hands, feet, and limb joints in many mammal ancestors, traits typically associated with sprawling postures among modern animals. It can also help explain why fossils of early mammal ancestors are frequently found in a squashed, spread-eagle pose – a pose more likely to be achieved with sprawled limbs, while modern placental and marsupial fossils are typically found lying on their sides. “It is very gratifying as a scientist, when one set of results can help illuminate other observations, moving us closer to a more comprehensive understanding,” Bishop said. Pierce, whose lab has studied the evolution of the mammalian body plan for nearly a decade, notes that these findings are consistent with patterns seen in other parts of the synapsid body, like the vertebral column. “The picture is emerging that the full complement of quintessentially therian traits was assembled over a complex and prolonged period, with the full suite attained relatively late in synapsid history,” she said. Conclusion: Complexity in Evolution Beyond mammals, the study suggests that some major evolutionary transitions, like the shift to an upright posture, were often complex and potentially influenced by chance events. For instance, the strong reversal in synapsid posture, back toward more sprawled poses, appears to coincide with the Permian-Triassic mass extinction—when 90% of life was wiped out. This extinction event led to other groups like the dinosaurs becoming the dominant animal groups on land, pushing synapsids back into the shadows. The researchers speculate that due to this “ecological marginalization,” the evolutionary trajectory of synapsids may have changed so much that it altered the way they moved. Whether this hypothesis turns out to be supported or not, understanding the evolution of mammal posture has long been a complex puzzle. Pierce emphasized how advances in computing power and digital modeling have provided scientists new perspectives to address these ancient mysteries. “Using these new techniques with ancient fossils allows us to have a better perspective of how these animals evolved, and that it wasn’t just this simple, linear evolutionary story,” she said. “It was really complicated and these animals were probably living and moving in their environments in ways that we hadn’t appreciated before. There was a lot happening and mammals today are really quite special.” Reference: “Late acquisition of erect hindlimb posture and function in the forerunners of therian mammals” by Peter J. Bishop and Stephanie E. Pierce, 25 October 2024, Science Advances. DOI: 10.1126/sciadv.adr2722

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